Rarity and abundance in a diverse African forest

Duncan Thomas

Abstract Kakamega forest is one of the remnants of the equatorial guineo rainforest in the Eastern fringes of Africa. It was perhaps cut-off from the Congo region in the early volcanic era when the Great Rift Valley was formed. The forest is known for its diversity of biotic species, and it is home to some of the rare plants in the East African region. It has some of the rare species of, birds, snakes, insects and primates.

Biodivers Conserv (2007) 16:2045–2074 DOI 10.1007/s10531-006-9065-2 ORIGINAL PAPER Rarity and abundance in a diverse African forest David Kenfack Æ Duncan W. Thomas Æ George Chuyong Æ Richard Condit Received: 10 April 2006 / Accepted: 15 May 2006 / Published online: 12 July 2006 Springer Science+Business Media B.V. 2006 Abstract We censused all trees ‡1 cm dbh in 50 ha of forest in Korup National Park, southwest Cameroon, in the central African coastal forest known for high diversity and endemism. The plot included 329,519 individuals and 493 species, but 128 of those taxa remain partially identified. Abundance varied over four orders of magnitude, from 1 individual per 50 ha (34 species) to Phyllobotryon spathulatum, with 26,741 trees; basal area varied over six orders of magnitude. Abundance patterns, both the percentage of rare species and the dominance of abundant species were similar to those from 50-ha plots censused the same way in Asia and Latin America. Rare species in the Korup plot were much less likely to be identified than common species: 42% of taxa with <10 individuals in the plot were identified to species, compared to 95% of the abundant taxa. Geographic ranges for all identified species were gleaned from the literature and online flora. Thirteen of the plot species are known only from Korup National Park (all discovered during the plot census), and 39 are restricted to the Nigeria–Cameroon coastal zone. Contrary to expectation, species with narrow geographic ranges were more abundant in the plot than average. The small number of narrow endemics (11% of the species), many locally abundant, mitigates short-term extinction risk, either from demographic stochasticity or habitat loss. D. Kenfack (&) Missouri Botanical Garden, 4500 Shaw Blvd., St, Louis, MO 63110, USA e-mail: david.kenfack@mobot.org D. W. Thomas Department of Forest Science, Oregon State University, Corvallis, OR 97331-2902, USA G. Chuyong Department of Life Sciences, University of Buea, P.O. Box 63, Buea, Cameroon R. Condit National Center for Ecological Analysis and Synthesis, 735 State St. Suite 300, Santa Barbara, CA 93101, USA R. Condit Smithsonian Tropical Research Institute, Unit 0948, APO AA 34002-0948, USA 123 2046 Biodivers Conserv (2007) 16:2045–2074 Keywords Korup Æ Cameroon Æ Tree abundance Æ Dominance Æ Rarity Æ Geographic range Introduction Rarity is central to tropical forest conservation. Diverse communities inevitably include large numbers of species which are seldom recorded: the singletons in many inventories. In typical forest plots of a single hectare or less, 30% of the tree species may be singletons (Pitman et al. 1999). To exacerbate the difficulties with our understanding of tropical communities, species identification is often problematic. Even experts leave many specimens as ‘morphospecies’—recognizable within a site, but not matched to named collections in herbaria. For these reasons—rarity and difficult taxonomy—quantitative information on the abundances and distributions of tropical organisms, not just trees, remains problematic, and conservation planning is hindered by lack of basic knowledge about which species are most endangered. This is probably true in Africa more than any other region. In addition, African forests can have unusual abundance patterns. One or a few species sometimes approach abundances observed in temperate forests (e.g. Sankovski and Pridnia 1995; Shaw et al. 2004), where a single dominant tree comprises more than half the forest (Makana 1998; Hart unpublished, 1990). This is known as monodominance, and although it is known in tropical forests elsewhere, it is most important in Africa (Marimon et al. 2001; Nascimento and Proctor 1997). Monodominance should go hand-in-hand with low diversity (Connell and Lowman 1989) and rarity, because the dominant trees force the scarce species to even lower abundance. Enhanced rarity exacerbates the problems of studying the forests. More exhaustive inventories are needed to document tree species abundances, and rarity might enhance extinction risk for many species. The coastal forests of Western Africa, especially Cameroon, are increasingly recognized as important for the conservation of forest diversity. They are the richest in plant species across Africa and repeatedly appear as a center of endemism for plants and for animals (Lovett et al. 2000; Linder 2001; Küper et al. 2004; Rodrigues et al. 2004; Burgess et al. 2005). Because conservation of species and communities depends on more than just species counts, we sought more detailed information about the forests of coastal Cameroon. As part of a global network of large census plots within the tropics, coordinated by the Center for Tropical Forest Sciences (CTFS) of the Smithsonian Institution, we initiated a large-scale and precise forest inventory, aimed at documenting the abundances of all tree species and testing hypotheses about factors that regulate diversity and species composition. The plot covers 50 ha in the Korup National Park, and all individual trees have been mapped and identified (Thomas et al. 2003; Chuyong et al. 2004). Along with a companion plot in the Democratic Republic of the Congo (Makana et al. 2004), these are the largest forest plot inventories in Africa, and provide quantitative assessments of abundance and rarity. Here we describe the floristic composition, structure, and physiognomy of the Korup plot. We answer several basic questions about tree abundance: is the forest monodominant, and how abundant are the dominant canopy tree species? Is the understory also dominated by one or a few species? At the opposite extreme, how 123 Biodivers Conserv (2007) 16:2045–2074 2047 many species are rare in 50 ha? We then consider whether the rare species at Korup have narrow geographic ranges, as predicted by macroecological theory (Brown 1995; Gaston 2003). Conservation planning often relies on the local abundance of species as well as their geographic ranges, and here we provide information for judging the conservation importance of southwest Cameroon (Rodrigues and Gaston 2000). Because a dozen other sites in the world now have comparable inventories (Condit et al. 2005), abundance patterns at Korup can be compared against the rest of the world, so that we can determine whether African forests differ fundamentally, as Richards (1973) once suggested. Materials and methods The park and forest The Korup plot (NW corner 503.86¢ N, 851.17¢ E) is located in southern Korup National Park (454¢ to 528¢ N latitude and 842¢ to 916¢ E longitude), near the coast of Cameroon and the Nigerian border. It is within a belt of evergreen forest extending from southeast Nigeria to the mouth of the Congo River (Fig. 1) that is called the Lower Guinea forest, containing one or more Pleistocene refugia (White 1979; Maley 1987). The belt is characterized by wet forested lowlands, often backed by mountain ranges, and is generally reported to be rich in species (Linder 2001). Letouzey (1968, 1985) described the southern part of the Korup National Park as Biafran coastal forest, rich in gregarious Fabaceae-Caesalpinioideae. Fig. 1 Map of tropical Africa showing the three major forest blocks, from west to east: Upper Guinea, Lower Guinea, Congo (or Congolian). Inset in lower left is the detailed map of southeast Nigeria and southwest Cameroon, with Korup National Park indicated 123 2048 Biodivers Conserv (2007) 16:2045–2074 The plot measures 1,000·5,00 m2 and is 150–240 m above sea level. The long axis of the plot crosses a permanent creek which is fed by a number of small, seasonal drainages (Fig. 2). One-third of the plot is a steep, rocky ridge, while the remaining two-thirds are flat terrain (Fig. 2). Soils of southern Korup are mostly non-hydric, skeletal, with a high sand content and low pH (Gartlan et al. 1986). Detailed soil mapping within the plot has been initiated but will be reported elsewhere. Census We first surveyed the 50 ha, placing permanent stakes precisely every 20 m with minimal damage to the vegetation (Condit 1998). Subsequently, all individuals ‡1 cm diameter-breast-height (dbh) were tagged with numbered aluminum tags, mapped, measured, and identified to morphospecies. Stem diameter was measured 1.3 m above the ground, with swollen or buttressed trees measured at a spot where the trunk was more regular; in these cases, the measurement spot was painted so future measurements could match. Any stem fork or branch <1.3 m above the ground was treated as a secondary stem and also measured. To map stems, ropes were strung between adjacent stakes (including diagonals) and temporary stakes were placed at 5-m intervals, then the position of each stem was marked by eye on maps covering 10·10 m2. Additional details are given in Condit (1998). Taxonomy and collection Most of the trees enumerated lacked flowers and fruits, so we had to rely on vegetative characters to segregate morphospecies: color, odor, and texture of bark slash; color of exudates from bark or leaves; leaf arrangement; petiole length; leaf shape; the number and form of secondary veins; and indumenta. Vouchers were collected from each morphospecies in the plot when first encountered. The first specimen was considered the ‘‘type’’ and was carefully described. Each morphospecies was subsequently collected at least five times (excepting those with <5 individuals) and compared side-by-side with the ‘‘type’’ until we were confident we could recognize Fig. 2 Topographic map of the Korup 50-ha plot, 1000 m · 500 m with 2 m contours. North and the highest point in the plot is to the left 123 Biodivers Conserv (2007) 16:2045–2074 2049 it. Flowers or fruits were eventually collected from most taxa. Matching and recognizing morphospecies is a matter of judgment, and our opinions changed through time, but two of us (DK, DT) spent 700 days over 3 years in the plot, viewed nearly every one of 330,000 individuals, and checked all morphospecies against reference collections at herbaria in Cameroon, the U.K., and the U.S. (Herbaria YA, SCA, K, MO) and against regional floras (Aubréville et al. 1963–2001, 1961–1991; Hutchinson et al. 1954–1972). We are reasonably confident in our current classification of individuals into 493 morphospecies, but a few will change when fertile material is studied. A few trees could not be sorted, mostly because they lacked leaves but appeared to be alive. Our family classification follows the Angiosperm Phylogeny Group (2003). Abundance and diversity calculations Because many inventories in tropical forest cover just 1 ha, we report abundance and basal area on a per hectare basis as well as for all 50 ha combined. The former is simply 0.02 times the latter, however, we can go further by estimating standard deviations by dividing the 1000·500 m2 into 50 non-overlapping 100 · 100 m2 and counting individuals (basal area) in each. Our counts of individuals do not include multiple stems per tree, but multiple stems are included in the basal area calculations. Unidentified trees were included in calculations of total plot basal area and density. For species richness of trees ‡1 cm dbh, we report the tally for all 50 ha and the mean per hectare, obtained by averaging the totals from 50 100·100-m2. The trees not identified to morphospecies were not included in diversity estimates. All calculations were repeated for trees ‡10 cm dbh. Height categories of species We classified all species into four growth-forms according to their estimated maximum height. Treelets and small trees include all species with adults generally less than 10 m tall; understory trees are those with adults 10–20 m tall; lower canopy species have heights 20–30 m; and upper canopy species are those often >30 m in height and emergent above the main canopy. Corresponding adult stem diameters were <10 cm, 10–30 cm, 30–60 cm, and >60 cm dbh, respectively. Information on the heights of the species came from field estimates in the plot supplemented with information from the literature, especially Aubréville et al. (1963–2001) and Hutchinson et al. (1954–72). Geographic range Distribution patterns of the Korup tree species were tallied relative to the major African phytochoria (White 1979, 1983), which are based on the three main blocks of moist tropical forest in Africa (Fig. 1). White’s large eastern block, or Congolian forest, falls largely in the two countries called Congo. His central block is the Lower Guinean forest, and covers the coastal belt from eastern Nigeria south through Gabon. The western block is Upper Guinea, mostly in Ivory Coast and Ghana. These three blocks are considered Pleistocene forest refugia (White 1983; Maley 1987), and Upper Guinea in the west is currently isolated from Lower Guinea by the Dahomey 123 2050 Biodivers Conserv (2007) 16:2045–2074 Gap, a low rainfall area where savanna reaches the Atlantic coast; Lower Guinea and the Congolian forest are presently contiguous. Tree distributions were obtained from the literature (Aubréville et al. 1963– 2001, 1961–1991; Hutchinson et al. 1954–1972), supplemented by the TROPICOS database (http://www.mobot.mobot.org/W3T/Search/vast.html). Ideally, we would have considered ranges quantitatively, but species distribution data from tropical Africa is too sparse to allow this. Instead, we assigned each species to one of seven categories: (1) pan-African, including all moist forest and extending into dry and montane forests around it; (2) Guineo–Congolian, including all three moist forest blocks; (3) Lower Guineo–Congolian, meaning the central and eastern blocks; (4) Upper and Lower Guinea, or the central and western blocks; (5) Lower Guinea only; (6) coastal Nigeria–Cameroon only; and (7) Korup National Park only. The only species in the last category are those we discovered in the 50ha plot. We tested the hypothesis that geographic range was associated with abundance within the 50-ha plot, and this required estimates of statistical confidence. For all species in the same category of geographic range, we calculated the median abundance per 50 ha. Confidence limits in those medians were estimated by a spatial form of bootstrapping, because highly aggregated species distributions do not justify standard statistics (Valencia et al. 2004). The plot was divided into 50 individual hectares (non-overlapping 100 · 100 m2), and these were resampled with replacement 1,000 times. For each sample, the median abundances were recalculated, and the 2.5th and 97.5th percentiles were used as confidence limits. Results Floristics and diversity A total of 493 morphospecies were recorded within the 50-ha plot, including 365 (71%) identified to species, 96 (20%) identified to genus, 29 (6%) identified to family, plus three not yet known at even the family level (Table 1). In addition, 680 individual trees have not been assigned a morphospecies. There were 245 genera and 62 families among the 493 species (Appendices 1–3). Among trees ‡10 cm dbh, there were 306 species, 184 genera, and 53 families (Table 1). So far, we have discovered 13 new species in the plot, and four of these have been described (Kenfack et al. 2004, 2006; Sonké et al. 2002; Gereau and Kenfack 2000). We anticipate more novel species among the 128 unnamed morphospecies and the 680 unassigned individuals. The family Rubiaceae was the richest in the plot, with 86 species in 40 genera; Fabaceae was next with 39 species in 25 genera. The traditional Euphorbiaceae had 37 species and 25 genera, but APG II (2003) divided this into the Phyllanthaceae (9 genera), the Euphorbiaceae (14 genera), and the Putranjivaceae (2 genera). The Annonaceae and Malvaceae (including the traditional Sterculiaceae and Tiliaceae) also had more than 20 species each in the plot, and the Annonaceae had 11 genera (Appendix 3). There were 233.1 species per ha among all individuals (Table 1), and 88.5 among trees ‡10 cm dbh. Fisher’s alpha for the entire plot was 56.9, and 62.8 for trees ‡10 cm dbh, but was lower for individual hectares (Table 1). 123 Biodivers Conserv (2007) 16:2045–2074 Table 1 Structure and diversity of the Korup 50-ha plot. N refers to the total number of, and BA the basal area. SD = standard deviation Mean N ha–1 SD N ha–1 Mean BA ha–1 SD BA ha–1 Mean species ha–1 SD species ha–1 Species (50 ha)–1 Mean Fisher’s ha–1 SD Fisher’s ha–1 Fisher’s (50 ha)–1 2051 ‡1 cm ‡10 cm ‡30 cm 6586.4 987.6 32.0 4.1 238 17.5 493 48.3 3.8 56.9 491.8 49.7 26.0 4.0 86.3 12.0 306 30.5 5.6 49.3 83.9 14.9 16.1 4.1 35.5 8.2 192 24.2 8.9 41.5 Abundance We recorded 329,319 trees with dbh ‡1 cm within the 50-ha plot, of which 24,591, or 7.5%, were ‡10 cm dbh; this amounts to 6,586 individuals ha–1, with 492 ha–1 above 10 cm dbh. The plot had 32.0 m2 ha–1 basal area, and 26.0 m2 ha–1 in trees ‡10 cm. Cola in the Malvaceae was the most abundant genus (Appendix 2), followed by Rinorea (Violaceae), Phyllobotryon (Salicaceae, formerly Flacourtiaceae), and Diospyros (Ebenaceae). The Malvaceae was the most abundant family, followed by Violaceae, Salicaceae, and the Euphorbiaceae (Appendix 3). Oubanguia was the dominant genus in basal area, almost entirely in one species, Oubanguia alata (Lecythidaceae, formerly Scytopetalaceae). Lecythidaceae was the dominant family in basal area, due mostly to Oubanguia alata, followed by Fabaceae, Malvaceae and Euphorbiaceae (Table 2). Several treelets were the most abundant species in the plot, with Phyllobotryon spathulatum first and three other Cola close behind (Table 2). Two canopy species, Oubanguia alata and Dichostemma glaucescens (Euphorbiaceae) were also among the top 10 species in abundance. Oubanguia and Dichostemma ranked first and third in total basal area, and Cola laterita ranked second, with low stem density but many large trees (Table 2). The largest diameters overall were mostly Lecomtedoxa klaineana (Sapotaceae), reaching 190 cm and including 13 of the 20 biggest trees; the single biggest tree was a 205-cm Erythrophleum ivorense (Fabaceae). At the other extreme, 221 of 493 species (45%) had a mean density of £1 tree ha–1, and 34 species were singletons in 50 ha. Considering only trees ‡10 cm dbh, there were 38 singletons in the 50 ha, and 239 species had density <1 ha–1 (78% of all species ‡10 cm). Basal area had an even greater range: 12 rare species had 2·10–6 m2 ha–1 (a single 1-cm sapling in 50 ha), while Oubanguia had 4.3 m2 ha–1. The distribution of basal area and abundance per species approached log-normal, though deviating slightly with an excess of rarity (Fig. 3). These highly skewed abundance distributions produce extremes in the way that a few species dominate: the 10 most abundant species (2% of the total) accounted for 42% of the individuals, while the 10 rarest species accounted for <0.1%. In basal area, the 10 dominants accounted for 41% of the forest, while the 10 rarest accounted for 0.0001%. Species rare in the plot were considerably less likely to be fully identified than abundant species. Of the 100 rarest species (£7 individuals), 47 were identified to species, in contrast to 89 fully identified out of the 100 most common species (>500 123 2052 123 Table 2 Dominant species in Korup 50-ha plot, in four diameter categories (‡10, ‡100, and ‡300 mm dbh), and in total basal area N ‡ 10 (rank) N ‡ 100 (rank) N ‡ 300 (rank) BA (rank) Phyllobotryon spathulatum (TL) (Lower Guinea) Cola semecarpophylla (TL) (SE Nigeria and SW Cameroon) Dichostemma glaucescens (C) (Lower Guinea–Congolian) Cola praeacuta (TL) (SE Nigeria and SW Cameroon) Oubanguia alata (C) (Lower Guinea) Cola sp. nov.2 (TL) (Korup) Cola sp. nov.3 (TL) (Korup) Diospyros preussii (C) (Lower Guinea) Angylocalyx oligophyllus (TL) (Upper and Lower Guinea) Rinorea lepidobotrys (TL) (Upper and Lower Guinea) Diospyros iturensis (U) (Lower Guinea–Congolian) Strombosia pustulata (C) (Guineo–Congolian) Drypetes staudtii (U) (SE Nigeria and SW Cameroon) Hymenostegia afzelii (C) (Upper and Lower Guinea) Diospyros gabunensis (C) (Upper and Lower Guinea) Protomegabaria stapfiana (C) (Upper and Lower Guinea) Cola rostrata (U) (Lower Guinea) Soyauxia gabonensis (U) (Lower Guinea) Klaineanthus gaboniae (C) (Lower Guinea) Strombosiopsis tetrandra (C) (Lower Guinea-Congolian) Lecomtedoxa klaineana (E) (Lower Guinea) Hypodaphnis zenkeri (C) (Lower Guinea) 534.6 490.4 345.0 309.4 298.4 247.3 164.7 147.1 115.9 109.8 95.9 82.4 82.0 80.5 78.0 67.3 65.7 63.6 40.6 35.3 6.1 6.0 0.1 0.4 45.5 9.1 82.1 0.0 0.2 0.0 0.0 1.9 10.2 15.7 21.7 13.8 17.1 18.7 18.4 10.0 9.7 6.3 4.2 3.1 0.0 0.0 1.9 0.0 18.1 0.0 0.0 0.0 0.0 0.0 0.0 2.4 0.1 1.6 0.9 7.0 0.2 0.4 2.8 2.4 2.9 1.6 0.2457 0.6660 1.5761 0.6971 4.3721 0.1378 0.1189 0.0668 0.0679 0.1910 0.2720 0.6881 0.5191 0.5540 0.5522 1.4642 0.5078 0.3345 0.6172 0.4609 1.9855 0.3205 (1) (2) (3) (4) (5) (6) (7) (8) (9) (10) (11) (15) (16) (17) (18) (24) (25) (28) (39) (42.5) (127) (129) (215) (121.5) (2) (14) (1) (369) (163.5) (369) (369) (42) (9) (7) (3) (8) (6) (4) (5) (10) (12) (17) (23) (31) (315) (315) (7) (315) (1) (315) (315) (315) (315) (315) (315) (5.5) (109.5) (8.5) (17) (2) (72.5) (46.5) (4) (5.5) (3) (8.5) (32) (7) (3) (5) (1) (53) (62) (86) (85) (40) (27) (6) (11) (9) (10) (4) (12) (18) (8) (15) (2) (21) N gives the total number of individuals per 50 ha ‡ 10 mm dbh, and BA the total basal area. Following each abundance category, the rank for that category is given in parentheses (rank 1 means the most abundant, 493 the least; in case of ties, all species involved were given the mean of the tied ranks, thus there are many ranks 344). A species is included if it was among the 10 most abundant species in any of the four categories. Examples: Hypodaphnis zenkeri is included because it ranked among the top 10 in abundance ‡300 mm dbh, although it ranked low in abundance ‡10 mm dbh. For each species, geographic range category and growth form (TL = treelet; U = understory; C = canopy; E = emergent) are listed Biodivers Conserv (2007) 16:2045–2074 Species 2053 40 30 0 10 20 Frequency 50 60 Biodivers Conserv (2007) 16:2045–2074 0.01 0.1 1 10 100 1e3 40 0 20 Frequency 60 80 N 1e-6 1e-5 1e-4 1e-3 1e-2 1e-1 1 10 BA Fig. 3 Abundance distributions for the 493 species in the Korup 50-ha plot. (A) Stem density ha–1. (B) Basal area ha–1. Abundance axis is logarithmic individuals). The estimated probability of identifying a species rose from 43% to 96% over the range of abundances in the plot (Fig. 4). Rarity in single hectares Per single hectare, there were 51.4 singletons ‡1 cm dbh, and 38.4 singletons ‡10 cm dbh (means of 50 individual hectares). These represent 22% and 44% of the total species richness ha–1 in the two size classes, respectively. Geographic range and abundance Of the species whose ranges were known, 123, or 33%, are known only from the Lower Guinean block of forest, from coastal Nigeria to Gabon (Table 3). Another 123 Biodivers Conserv (2007) 16:2045–2074 0.81 0.40 .6 species .2 probability of identification .0 2054 0.00 family or less 1e0 1e1 1e2 1e3 1e4 individuals in plot Fig. 4 Probability species are identified as a function of their abundance in 50 ha. Points are observed fraction of species in log-10 bins (1–9 individuals per 50 ha, 10–99, etc.). Curves are fitted logistic regressions based on all individual species, with binomial error. Upper curve and points give probability that morphospecies are fully identified to species; lower curve is the converse probability, not being identified to species 39 species have even narrower ranges, found only in the small area near the Cameroon–Nigeria border; this includes the 13 species discovered during the plot census, which are (for now) known only from Korup National Park. Thus, 162 species have ranges <300,000 km2, of which 39 species span <80,000 km2. The other 202 species have wider ranges, including 82 found in all three tropical African forests blocks. The species with narrow geographic ranges tended to be more common in the plot than widespread species. Only the 13 new species from Korup deviated from this trend; otherwise, the species in narrower range categories had median abundances well above the plot’s overall median (Table 3). Conversely, species abundant in the Table 3 Geographic range and abundance of species found in the Korup 50-ha plot. N is the number of individuals ‡10 mm dbh; CI gives the bootstrap 95% confidence limits on median abundance 123 Range Land No. Abundance (CI) area species median (103 km2) N ha–1 Widespread 1,800 in Africa Guineo–Congolian 1,750 Lower 1,600 Guinea–Congolian Upper and Lower 300 Guinea Lower Guinea 150 endemic 23 SE Nigeria and SW Cameroon Korup 1.29 Unknown All species 22 1.16 (0.93–1.38) 60 44 0.93 1.67 (0.77–1.16) (1.37–1.91) 76 1.74 (1.55–2.20) 123 3.34 (2.90–3.96) 26 8.19 (5.78–9.57) 13 129 493 1.02 0.28 1.42 (0.70–1.66) (0.20–0.34) (1.20–1.50) Biodivers Conserv (2007) 16:2045–2074 2055 plot had narrow ranges. For example, of the 10 most abundant species, 4 are narrow endemics, all treelets in the genus Cola, and 3 others are known only from Lower Guinea (Table 2). The comparison of medians, though, conceals wide variation in abundance: all geographic range categories included species from rare to common (Fig. 5). Life form Of the 493 species in the plot, 222 were treelets, mostly <10 m tall, including many of the most abundant species in the forest. Seven of the ten dominant species in terms of stem density were treelets, four of them in the genus Cola (Table 2). Another 85 species in the plot were understory trees <20 m tall, seldom reaching the canopy. All told, 307 species were thus largely confined to the understory, while 112 were main canopy species and 74 were emergent trees. Several of the treelet species have an unusual form, with a single unbranched stem bearing a terminal rosette of large leaves. The two most abundant species in the plot fall in this category (Phyllobotryon spathulatum and Cola semecarpophylla), plus several other Cola’s, Alexis cauliflora, and Crotonogynopsis sp. nov. The rosettes form wide, shallow funnels that collect leaf litter, which decomposes in situ to form a ball of humus over the stem apex. Discussion We discovered in a census of half a square kilometer nearly 500 tree species, and a quarter of these remain unidentified. To date, we know that 13 of the species were Phyllobotryon Dichostemma 100 Angylocalyx - 10 1 Cola sp2 Cola sp3 Diospyros Rinorea - - - - 0.1 abundance Cola se. Cola pr. Oubanguia - - Albizzia f. Leptaulus h. Coffea l. Garcinia p. Pauridiantha v. Allophylus h. Cola m. Draceana g. Spathodea c. Irivngia s. Rothmannia l. Engomagoma g. Manlikara l. Africa Gui-Cong L Gui-Cong U&L Gui L Gui Nig-Cam Korup range Fig. 5 Abundance of each species as a function of geographic range. The ranges are categorical, ordered from the most widespread (Africa) to the narrowest (Korup NP only); the position along the axis does not reflect range size quantitatively. Each point shows the abundance, on a log-scale, of a single species. Several of the most and least abundant species are named 123 2056 Biodivers Conserv (2007) 16:2045–2074 previously unknown, and we expect this number to increase a lot. Most of the 128 unidentified species, though, are probably already described and many of them belong to difficult genera with many species and poorly-documented intra-specific variation. Full identification of these difficult taxa is slow, requiring the careful examination of material in several European herbaria. Pushing the minimum diameter limit down to 1 cm, in contrast to the 10-cm limit used in most African tree inventories, uncovered a high density and diversity in the understory. Over 300 of the species in the plot typically reproduce in the forest understory, at least according to our preliminary observations on diameter distributions. Many of these did grow beyond 10 cm dbh, but 187 did not: our census would have included only 306 species had we not included the smaller trees. The understory is dominated by an odd group of litter-trapping treelets, several in the diverse African genus Cola. The size of the plot also allows us to document rarity. The Korup abundance distribution has a mode well above the rarest species, nevertheless, there is a long tail of uncommon species. Abundance varied over four orders of magnitude, from singletons in 50 ha to Phyllobotryon spathulatum, with nearly 27,000 trees; basal area varied by six orders of magnitude, from species with single individuals of 1 cm dbh to Oubanguia alata, with 4 m2 ha–1. Surprisingly, though, Korup abundances are no different from other tropical forests, indeed all large plots have very similar ranges of abundance and abundance distributions (Condit et al. 2005). At Barro Colorado in Panama, 45% of 301 species had <1 tree ha–1, exactly the figure at Korup, and at Pasoh in Malaysia, it was 34%. The most abundant species at Korup, including individuals above 1 cm dbh, had 535 trees ha–1, compared to 808 ha–1 at Barro Colorado and 179 ha–1 at Pasoh (Condit et al. 1996a; Manokaran et al. 1992). Among trees above 10-cm dbh, though, Oubanguia at Korup was strikingly abundant, with 82 trees ha–1, compared to Trichilia tuberculata (Meliaceae) in Panama, with 35 ha–1 and Xerospermum noronhainum (Sapindaceae) at Pasoh, with 11 ha–1. A parallel large inventory in monodominant forest in the Congo, however, found abundances far different from those at Korup (Makana et al. 1998, 2004; Condit et al. 2005). The dominant species above 10 cm dbh, Gilbertiodendron dewevrei (Fabaceae), had 180 trees ha–1, and in the understory, Scaphopetalum dewevrei (Malvaceae) had a remarkable 3,067 ha–1 trees above 1 cm dbh. At the rare end, 72% of the species there had <1 tree ha–1. Although Oubanguia alata is very abundant at Korup, the plot in Cameroon does not resemble the monodominant stand in the Congo. Stands of gregarious legumes occur elsewhere in Korup (Newbery and Gartlan 1996; Newbery et al. 2004) and these are not monodominant either. We therefore conclude that the Korup forest is not monodominant, and is more typical of mixed-species tropical forests worldwide. If Korup is typical of other tropical forests in terms of rarity, this should not deflect from the importance of rarity: all tropical forests have many rare species and these are the species with the greatest extinction risks. Unfortunately, they are also the most difficult to identify—among those species at Korup for which we had fewer than 10 individuals to examine, most are unidentified. We are thus missing information on geographic range for most of the rare species, which can skew our conclusions about rarity and extinction risk (Ruokolainen et al. 2002). Linder (2001) and Burgess et al. (2005) considered ‘red-list’ species to be those with ranges <2 square degrees of latitude, about 20,000 km2 following the IUCN threatened list. This is approximately the area of southeast Nigeria and southwest 123 Biodivers Conserv (2007) 16:2045–2074 2057 Cameroon that we consider an area of narrow endemism, to which 39 of the plot species are restricted. Species abundances are also relevant to extinction risk due to demographic stochasticity, and the plot data allow us to explore this. Of the 13 species restricted to Korup National Park, 9 have ‡47 individuals, and thus have estimated populations throughout the park of several thousand trees. Overall, it does not seem likely that demographic stochasticity is a threat to plot species. What about those species so rare we missed them? Based on evidence from Panama and Malaysia (Condit et al. 1996b) and our field work around the park, we estimate that the 50-ha plot includes on the order of half the tree species occurring in the park. Some of these may have populations rare enough that extinction due to demographic fluctuations is a plausible risk; but if these unobserved species follow the patterns observed in the plot, there would still be very few species at risk. On the other hand, the 39 species restricted to 20,000 km2 near Korup may be at some risk of extinction due to habitat loss. If most of the forest in this area were cleared, and much is already cultivated, even the abundant among these could approach extinction. Burgess et al. (2005) found 26% of the species they analyzed had ranges below the 20,000-km2 limit, compared to our estimate of 11% of the tree species. Our lower figure may be because trees have wider ranges than other life forms, since the Burgess analysis included a greater variety of angiosperm taxa. On the other hand, in forests around the Panama Canal, we found 1.1% of 630 tree species had highly restricted distributions, limited to the 3,000 km2 Panama Canal watershed (Condit et al. 2001), an area about twice the size of the Korup National Park. At total of 9.6% were endemic to Panama and Costa Rica, which at 127,000 km2 approaches the size of the Lower Guinea forest block. The Korup plot includes 162 tree species restricted to Lower Guinea, or 44% of the total, suggesting that coastal central Africa indeed has considerably narrower tree ranges than Central America. Our knowledge of rare tropical trees remains scanty, however, they are hard to find and hard to identify. Fully documenting the flora of tropical Africa requires considerably more work—thorough field inventories at many more sites. We hazard an estimate that 10%–20% of the flora remains undiscovered, and 25%–40% remains so poorly known that geographic ranges are just crude approximations. Large and precise inventories uncover new species and reveal a window into how many remain unknown. Acknowledgments Funding for the fieldwork in Cameroon was provided by the International Cooperative Biodiversity Groups (a consortium of the U.S. National Institutes of Health, the U.S. National Science Foundation, and the U.S. Department of Agriculture), with supplemental funding by the Central Africa Regional Program for the Environment (a program of the U.S. Agency for International Development), and the Celerity Foundation at the Peninsula Community Foundation. Permission to conduct the field program in Cameroon was provided by the Ministry of Environment and Forests and the Ministry of Scientific and Technical Research. Data analyses were supported by U.S. National Science Foundation grant #009011 to the Center for Tropical Forest Science of the Smithsonian Tropical Research Institute, a global network of large-scale demographic plots. 123 2058 Biodivers Conserv (2007) 16:2045–2074 Appendices Appendix 1 Species of the Korup 50-ha plot Genus Species Family Code N BA Form Range Afzelia Afzelia Aidia Albizia Allexis Allophylus Allophylus Allophylus Alsodeiopsis Alstonia Amphimas Angylocalyx Angylocalyx Anisophyllea Anisophyllea Anisophyllea Annickia Anthocleista Anthocleista Anthonotha Anthonotha Anthonotha Anthonotha Antidesma Antidesma Antrocaryon Aoranthe Aporrhiza Araliopsis Asystasia Aulacocalyx Aulacocalyx Aulacocalyx Baikiaea Baillonella Baphia Baphia Baphia Barteria Beilschmiedia Beilschmiedia Beilschmiedia Beilschmiedia Beilschmiedia Beilschmiedia Beilschmiedia Belonophora Belonophora Berlinia Berlinia Berlinia Bersama Bertiera bella bipindensis genipiflora ferruginea cauliflora africanus hirtellus megaphyllus weissenborniana boonei ferrugineus oligophyllus pynaertii meniaudii purpurascens sororia chlorantha schweinfurthii vogelii fragrans lamprophylla macrophylla sp. laciniatum vogelianum micraster cladantha sp. soyauxii macrophylla caudata jasminiflora talbotii insignis toxisperma capparidifolia laurifolia sp. fistulosa acuta jacques-felixii sp. sp. 1 sp. 2 sp. 3 sp. 4 talbotii wernhamii auriculata craibiana hollandii sp. laxa Fabaceae Fabaceae Rubiaceae Fabaceae Violaceae Sapindaceae Sapindaceae Sapindaceae Icacinaceae Apocynaceae Fabaceae Fabaceae Fabaceae Anisophylleaceae Anisophylleaceae Anisophylleaceae Annonaceae Gentianaceae Gentianaceae Fabaceae Fabaceae Fabaceae Fabaceae Phyllanthaceae Phyllanthaceae Anacardiaceae Rubiaceae Sapindaceae Rutaceae Acanthaceae Rubiaceae Rubiaceae Rubiaceae Fabaceae Sapotaceae Fabaceae Fabaceae Fabaceae Passifloraceae Lauraceae Lauraceae Lauraceae Lauraceae Lauraceae Lauraceae Lauraceae Rubiaceae Rubiaceae Fabaceae Fabaceae Fabaceae Melianthaceae Rubiaceae AFZI AFBI RUBT2 ALFE ALCA ALAF ALLO KLGH ERYT ALBO SAPS ANTA FAB3 ANME ANPU ANSO ENCH ANSC ANNO ANFR ANLA ANMA ANTH ANTI ANVO ANTR POCL SAP5 ARSO ACAN ROT2 AUCA AUTA BAIN BATO BAPI BAP2 BAPS BAFI BEI4 BEI3 BEIH BEI2 BEI6 BEIP BEIS BELI BEWE BEAU BECR BEHO WSAP BELA 82 3 33 1 883 3 1 122 16 66 22 5801 57 13 52 157 707 19 40 247 89 76 7 18 706 8 79 171 168 8 631 258 1846 20 4 1769 1069 51 79 324 365 1 1352 372 181 475 1467 1081 726 3 101 9 54 5.155 0.7377 0.2303 0.509 1.2924 7e-04 1e-04 0.0261 0.0022 14.8402 0.8963 3.3937 0.5717 0.6697 1.5137 2.4597 8.3172 1.5124 2.1206 8.8175 0.4126 1.9556 0.8031 0.0804 2.0852 0.0091 0.5942 0.6366 3.1839 0.0011 0.3988 0.4781 2.3689 0.0588 1.0854 12.8059 4.3992 0.2982 0.3336 0.5498 0.449 0.0665 1.7262 4.2152 0.1994 2.2991 0.495 0.2021 5.053 0.0607 0.8533 0.0048 0.0062 Emergent Emergent Understory Canopy Treelet Treelet Treelet Treelet Treelet Emergent Emergent Treelet Canopy Canopy Canopy Canopy Canopy Canopy Canopy Emergent Understory Understory Emergent Treelet Understory Emergent Understory Understory Emergent Treelet Treelet Treelet Understory Canopy Emergent Canopy Canopy Understory Understory Canopy Canopy Understory Treelet Canopy Understory Understory Treelet Treelet Canopy Understory Emergent Treelet Treelet L Gui-Cong L Gui-Cong U&L Gui Gui-Cong L Gui Africa L Gui Nig-Cam L Gui Gui-Cong L Gui U&L Gui L Gui-Cong U&L Gui L Gui L Gui L Gui-Cong Gui-Cong Gui-Cong U&L Gui L Gui Gui-Cong . U&L Gui L Gui-Cong Gui-Cong L Gui-Cong . L Gui L Gui L Gui L Gui-Cong L Gui Gui-Cong L Gui L Gui U&L Gui . L Gui-Cong L Gui-Cong L Gui . . . . . Nig-Cam Nig-Cam U&L Gui L Gui Nig-Cam . L Gui 123 Biodivers Conserv (2007) 16:2045–2074 2059 Appendix 1 continued Genus Species Family Code N BA Form Range Bertiera racemosa Rubiaceae RUB5 671 0.1268 Treelet Gui-Cong Blighia sp. Sapindaceae COLL 20 0.4155 Emergent . Brenania brieyi Rubiaceae OPLA 10 0.0057 Emergent L Gui Bridelia micrantha Phyllanthaceae BRMI 45 0.4337 Canopy Africa Bridelia sp. Phyllanthaceae BRID 14 0.4054 Canopy . Caloncoba glauca Salicaceae CAGL 88 6.6273 Canopy L Gui Calpocalyx dinklagei Fabaceae CADI 3066 7.7904 Understory L Gui Calycosiphonia spathicalyx Rubiaceae BEL2 112 0.1034 Treelet L Gui-Cong Camptostylus mannii Achariaceae CAMP 143 0.3376 Understory L Gui Campylospermum calanthum Ochnaceae OURA 498 0.5449 Treelet L Gui Campylospermum flavum Ochnaceae OUR2 221 0.0351 Treelet U&L Gui Campylospermum mannii Ochnaceae OUEL 222 0.0573 Treelet Nig-Cam Campylospermum sulcatum Ochnaceae OUR6 91 0.0132 Treelet U&L Gui Canthium sp. 1 Rubiaceae CABL 1 1e-04 Treelet . Canthium sp. 2 Rubiaceae CAN4 33 0.1148 Canopy . Canthium sp. 3 Rubiaceae CANI 145 0.2011 Understory . Carapa dinklagei Meliaceae CABM 18 1.0477 Canopy L Gui Carapa parvifolia Meliaceae CAPR 782 7.1042 Canopy L Gui Carpolobia lutea Polygalaceae CALU 485 0.1245 Treelet U&L Gui Casearia barteri Salicaceae WHOM 80 0.3792 Understory L Gui Cassipourea korupensis Rhizophoraceae RUDR 47 0.1495 Understory Korup Ceiba pentandra Malvaceae CEPE 2 0.0114 Emergent Africa Cephaelis sp. Rubiaceae CEPH 10 0.004 Treelet . Chrysobalanus icaco Chrysobalanaceae WIRG 62 0.4534 Canopy Africa Chrysophyllum delevoyi Sapotaceae GAAF 33 0.3322 Canopy U&L Gui Chrysophyllum sp. nov. Sapotaceae SAPO 160 0.3549 Understory Korup Chytranthus sp. 1 Sapindaceae CHY1 1 3e-04 Treelet . Chytranthus sp. 2 Sapindaceae CHY2 47 0.0105 Treelet . Chytranthus sp.3 Sapindaceae CHY4 89 0.0347 Treelet . Chytranthus talbotii Sapindaceae CHTA 345 0.4915 Understory U&L Gui Cleistanthus letouzeyi Phyllanthaceae WDAS 43 0.2203 Understory L Gui Cleistopholis patens Annonaceae CLPA 38 0.7995 Canopy Gui-Cong Cleistopholis staudtii Annonaceae CLSP 11 1.5895 Canopy L Gui Coelocaryon preussii Myristicaceae COPR 87 5.3102 Emergent L Gui-Cong Coffea liberica Rubiaceae COSS 1 2e-04 Treelet Gui-Cong Cola acuminata Malvaceae CONI 189 2.8933 Understory Gui-Cong Cola altissima Malvaceae COAL 4 0.0019 Canopy L Gui Cola cauliflora Malvaceae COCA 4272 4.8927 Treelet L Gui Cola chlamydantha Malvaceae COCH 191 0.3249 Understory U&L Gui Cola digitata Malvaceae CODI 414 0.1433 Treelet U&L Gui Cola flaviflora Malvaceae COTL 23 0.016 Treelet L Gui Cola gabonensis Malvaceae COGA 4 5e-04 Treelet L Gui Cola heterophylla Malvaceae COLS 47 0.0102 Treelet U&L Gui Cola lateritia Malvaceae COLA 465 16.2496 Emergent Gui-Cong Cola lepidota Malvaceae COLE 285 1.5807 Understory L Gui Cola marsupium Malvaceae COMA 1 0.002 Treelet L Gui Cola megalophylla Malvaceae COME 20 0.8221 Emergent Nig-Cam Cola pachycarpa Malvaceae COPA 27 0.1311 Understory L Gui Cola praeacuta Malvaceae COL2 15498 34.8528 Treelet Nig-Cam Cola rostrata Malvaceae CORO 3288 25.3913 Understory L Gui Cola semecarpophylla Malvaceae COSE 24535 33.3014 Treelet Nig-Cam Cola sp. 2 Malvaceae COBE 1 3e-04 Treelet . Cola sp. 5 Malvaceae COLSL 2 2e-04 Treelet . Cola sp. 6 Malvaceae COMT 4 0.002 Treelet . Cola sp. nov. 2 Malvaceae COAT 12396 6.8912 Treelet Korup 123 2060 Biodivers Conserv (2007) 16:2045–2074 Appendix 1 continued Genus Species Family Code N BA Form Range Cola sp. nov. 3 Malvaceae OCTI 8244 5.9463 Treelet Korup Cola suboppositifolia Malvaceae CONS 358 0.14 Treelet Nig-Cam Cola verticillata Malvaceae COVE 87 4.2119 Emergent U&L Gui Copaifera mildbraedii Fabaceae CAE3 8 0.1553 Emergent L Gui-Cong Cordia sp. Boraginaceae SIFB 5 0.0482 Understory . Coula edulis Olacaceae COED 8 0.3554 Canopy U&L Gui Crateranthus talbotii Lecythidaceae CRTA 18 0.0246 Understory Nig-Cam Craterispermum aristatum Rubiaceae CRAR 2625 2.7173 Treelet Nig-Cam Craterispermum caudatum Rubiaceae RUGA 59 0.0306 Treelet U&L Gui Croton longiracemosus Euphorbiaceae NEGL 112 3.9922 Canopy . Crotonogyne strigosa Euphorbiaceae CRST 3888 1.2752 Treelet L Gui Crotonogynopsis sp. nov. Euphorbiaceae ALEX 2158 0.9347 Treelet Korup Crudia sp. Fabaceae CRUD 1 0.2489 Emergent . Dacryodes edulis Burseraceae DAED 75 5.0607 Canopy L Gui Dacryodes klaineana Burseraceae DAKL 114 2.2306 Canopy U&L Gui Dactyladenia pallescens Chrysobalanaceae ACI2 155 0.5435 Understory L Gui Dactyladenia staudtii Chrysobalanaceae ACST 75 0.1607 Understory L Gui Dasylepis blackii Achariaceae DABL 2144 7.2305 Understory L Gui Deinbollia maxima Sapindaceae CHY3 501 0.2791 Treelet U&L Gui Deinbollia pychnophylla Sapindaceae COSL 109 0.7361 Understory L Gui Deinbollia unijuga Sapindaceae DEUN 6 7e-04 Treelet Korup Desbordesia glaucescens Irvingiaceae DEGL 85 25.0731 Emergent L Gui-Cong Dialium pachyphyllum Fabaceae DIAL 353 3.5118 Canopy L Gui Dialium sp. Fabaceae DIA3 66 2.4708 Canopy . Dichaetanthera africana Melastomataceae DIAF 3 0.0213 Treelet Gui-Cong Dichostemma glaucescens Euphorbiaceae DIGL 17255 78.804 Canopy L Gui-Cong Dicranolepis disticha Thymelaeaceae DIDI 9 0.0017 Treelet U&L Gui Dicranolepis sp. Thymelaeaceae DICR 2 0.1495 Treelet . Didymosalpinx sp. Rubiaceae RUBT 470 0.5067 Treelet . Dinophora spenneroides Melastomataceae DINO 2 2e-04 Treelet Gui-Cong Diogoa zenkeri Olacaceae DIOG 3220 12.4436 Canopy L Gui-Cong Diospyros bipindensis Ebenaceae DIBI 3 0.0095 Understory L Gui-Cong Diospyros gabunensis Ebenaceae DIGA 3899 27.6102 Canopy U&L Gui Diospyros gracilescens Ebenaceae DICI 163 2.5099 Emergent L Gui Diospyros hoyleana Ebenaceae DIHO 281 0.6836 Understory Gui-Cong Diospyros iturensis Ebenaceae DIIT 4795 13.6023 Understory L Gui-Cong Diospyros mannii Ebenaceae DIMA 87 0.3206 Emergent U&L Gui Diospyros obliquifolia Ebenaceae DIOB 729 0.3901 Treelet L Gui Diospyros physocalycina Ebenaceae DIO2 9 0.0082 Treelet L Gui Diospyros preussii Ebenaceae DIPR 7359 3.3379 Treelet L Gui Diospyros pseudomespilus Ebenaceae DIPS 1039 3.3276 Canopy Nig-Cam Diospyros sp. 1 Ebenaceae DIBM 8 0.4093 Canopy . Diospyros sp. 2 Ebenaceae DIOY 111 0.0846 Treelet . Diospyros sp. 3 Ebenaceae NDIO 5 0.1415 Canopy Gui-Cong Diospyros zenkeri Ebenaceae DIZE 655 4.8005 Understory L Gui Discoclaoxylon hexandrum Euphorbiaceae CLHE 37 0.2272 Canopy Gui-Cong Discoglypremna caloneura Euphorbiaceae DICA 167 6.7191 Emergent Gui-Cong Dorstenia turbinata Moraceae DORS 10 0.001 Treelet U&L Gui Dracaena bicolor Ruscaceae DRAI 171 0.088 Treelet U&L Gui Dracaena cf.goldieana Ruscaceae DRAT 1 1e-04 Treelet Korup Dracaena laxissima Ruscaceae DRAJ 54 0.0081 Treelet Africa Dracaena sp. Ruscaceae DRA2 12 0.0062 Treelet . Dracaena surculosa Ruscaceae DRA3 4 5e-04 Treelet U&L Gui Drypetes ivorensis Putrangivaceae DRYS 113 0.1428 Treelet U&L Gui Drypetes laciniata Putrangivaceae DRLA 249 0.138 Treelet L Gui Drypetes molunduana Putrangivaceae DRY3 26 0.0091 Treelet Nig-Cam 123 Biodivers Conserv (2007) 16:2045–2074 2061 Appendix 1 continued Genus Species Family Code N BA Form Range Drypetes principum Putrangivaceae DRSI2 3043 1.7722 Treelet U&L Gui Drypetes sp. Putrangivaceae CAR2 47 0.2281 Understory . Drypetes staudtii Putrangivaceae DRST 4101 25.9534 Understory Nig-Cam Duboscia macrocarpa Malvaceae DUMA 2 0.0012 Canopy L Gui-Cong Endodesmia calophylloides Clusiaceae ENCA 270 3.4477 Emergent L Gui Englerophytum sp. nov. Sapotaceae ENGI 652 2.4754 Understory Korup Engomegoma gordonii Olacaceae WCOU 1 0.5555 Canopy L Gui Entandrophragma cylindricum Meliaceae ENTI 44 0.0585 Emergent Gui-Cong Eriocoelum sp. Sapindaceae ERIO 76 1.5052 Canopy . Erismadelphus exsul Vochysiaceae EREX 251 13.7765 Emergent L Gui Erythrina sp. Fabaceae ERAD 8 0.1152 Canopy . Erythrophleum ivorense Fabaceae ERIV 79 24.5632 Emergent U&L Gui Erythroxylum mannii Rhizophoraceae ALFB 21 1.3316 Emergent U&L Gui Euclinia longiflora Rubiaceae EULO 131 0.0872 Treelet Gui-Cong Eugenia callophyloides Myrtaceae EUG2 30 0.0273 Treelet U&L Gui Eugenia fernandopoana Myrtaceae EUGT 24 0.0885 Understory L Gui Eugenia talbotii Myrtaceae EUGI 285 0.0433 Treelet Nig-Cam Eurypetalum unijugum Fabaceae EUUN 7 0.3198 Understory L Gui Fagara macrophylla Rutaceae ZAMA 109 2.7442 Canopy L Gui-Cong Ficus sp. Moraceae FICU 3 0.0773 Treelet . Funtumia elastica Apocynaceae FUEL 11 0.2239 Canopy Gui-Cong Gaertnera bieleri Rubiaceae GABI 714 0.1575 Treelet L Gui-Cong Gaertnera letouzeyi Rubiaceae GAET 7 0.001 Treelet Korup Garcinia conrauana Clusiaceae GACO 1287 11.5199 Understory L Gui Garcinia Guietoides Clusiaceae GAR2 1141 1.5443 Understory U&L Gui Garcinia kola Clusiaceae GAKO 20 0.0935 Canopy Gui-Cong Garcinia mannii Clusiaceae GAMA 158 0.6207 Understory L Gui Garcinia ovalifolia Clusiaceae GAOV 151 0.2874 Understory Gui-Cong Garcinia polyandra Clusiaceae GATB 1 2e-04 Treelet Gui-Cong Garcinia smeathmannii Clusiaceae GAR4 38 0.2925 Understory Gui-Cong Garcinia sp.1 Clusiaceae GAR3 29 0.0086 Treelet . Garcinia sp. 2 Clusiaceae GAR5 2 4e-04 Treelet . Garcinia sp. 3 Clusiaceae NGAR 1 1e-04 Treelet . Gilbertiodendron demonstrans Fabaceae GIDE 288 2.0869 Understory L Gui Glossocalyx brevipes Monimiaceae GLBR 1070 0.8259 Treelet L Gui Glyphaea sp. Malvaceae HGRE 14 0.0091 Treelet . Guarea glomerulata Meliaceae GUGL 17 0.0064 Treelet L Gui Guarea sp. Meliaceae GUA2 16 0.005 Treelet . Guarea thompsonii Meliaceae GUAR 116 7.8413 Emergent Gui-Cong Hallea ledermannii Rubiaceae MICI 69 5.5641 Emergent Gui-Cong Heckeldora staudtii Meliaceae SMEL 6 8e-04 Treelet L Gui Heinsia crinita Rubiaceae HECR 103 0.359 Treelet Africa Heisteria parvifolia Olacaceae HEPA 1174 0.1943 Treelet U&L Gui Homalium africanum Salicaceae HOMO 45 3.0684 Canopy U&L Gui Homalium africanum Salicaceae HOSA 88 0.4298 Understory L Gui Homalium letestui Salicaceae HOM2 618 2.5702 Emergent U&L Gui Homalium longistylum Salicaceae HOLO 535 8.9453 Emergent Africa Homalium sp. Salicaceae HOM4 1 0.0028 Understory . Hoplestigma kleineanum Hoplestigmataceae ALTB 19 0.3611 Canopy U&L Gui Hunteria umbellata Apocynaceae HUUM 874 8.462 Canopy Gui-Cong Hylodendron gabunense Fabaceae HYGA 13 1.9989 Emergent L Gui-Cong Hymenostegia afzelii Fabaceae HYAF 4025 27.6981 Canopy U&L Gui Hymenostegia bakeriana Fabaceae HYBA 203 6.4259 Emergent Nig-Cam Hymenostegia sp. nov. Fabaceae CAE2 51 2.4632 Emergent Korup Hypodaphnis zenkeri Lauraceae HYZE 299 16.026 Canopy L Gui Irvingia gabonensis Irvingiaceae IRGA 134 10.0348 Emergent U&L Gui Irvingia grandifolia Irvingiaceae IRGR 15 0.5895 Emergent L Gui-Cong 123 2062 Biodivers Conserv (2007) 16:2045–2074 Appendix 1 continued Genus Species Family Code N BA Form Range Irvingia Isolona Isolona Ixora Ixora Jollydora Jollydora Keayodendron Kigelia Klaineanthus Klainedoxa Klainedoxa Korupodendron Laccodiscus Laccodiscus Lannea Lasianthera Lasianthus Lasiodiscus Lecomtedoxa Lepidobotrys Leptaulus Leptaulus Leptonychia Leptonychia Licania Lophira Macaranga Maesobotrya Maesobotrya Maesobotrya Maesopsis Magnistipula Magnistipula Magnistipula Mammea Manilkara Maprounea Maranthes Mareya Mareyopsis Margaritaria Massularia Memecylon Memecylon Memecylon Memecylon Memecylon Memecylon Microcos Microdesmis Monodora Morelia Morinda Mostuea Musanga Mussaenda smithii campanulata sp. hippoperifera nematopoda duparquetiana glandulosa bridelioides africana gaboniae gabonensis trillesii songweanum ferrugineus pseudostipulari sp. africana batangensis sp. klaineana staudtii daphnoides holstii echinocarpa pallida elaeosperma alata monandra barteri dusenii staudtii eminii cuneatifolia glaberrima sp. africana lososiana membranacea sp. micrantha longifolia discoidea acuminata afzelii englerianum lateriflorum laurentii sp. zenkeri coriacea puberula brevipes senegalensis lucida brunonis cecropioides sp. Irvingiaceae Annonaceae Annonaceae Rubiaceae Rubiaceae Connaraceae Connaraceae Euphorbiaceae Bignoniaceae Euphorbiaceae Irvingiaceae Irvingiaceae Vochysiaceae Sapindaceae Sapindaceae Anacardiaceae Icacinaceae Rubiaceae Rhamnaceae Sapotaceae Lepidobotryaceae Icacinaceae Icacinaceae Malvaceae Malvaceae Chrysobalanaceae Ochnaceae Euphorbiaceae Phyllanthaceae Phyllanthaceae Phyllanthaceae Rhamnaceae Chrysobalanaceae Chrysobalanaceae Chrysobalanaceae Clusiaceae Sapotaceae Euphorbiaceae Chrysobalanaceae Euphorbiaceae Euphorbiaceae Phyllanthaceae Rubiaceae Memecylaceae Memecylaceae Memecylaceae Memecylaceae Memecylaceae Memecylaceae Malvaceae Pandaceae Annonaceae Rubiaceae Rubiaceae Gelsemiaceae Urticaceae Rubiaceae IRVI MONO WMON IXHI RUBI JOLY JOL2 KEBR KIAF KLGI KLGA KLTR PTHY LAFE LACC LANN LAAF LABA LASI LEKL PAPI LEDA LEPT LEEC LEPA LIEL LOAL MAMO MABA MADU MAST MAEM MAGUI MAGL CHR2 MAAF ENG2 MAME CHRY MAMI MALO MADI MAAC MEM4 MEML MEMT3 MEMT WMEM WWAR GRCO MIPU MOBI RUDO MOLU MOBR MUCE MUSA 1 256 86 552 1446 585 583 2 8 2032 8 7 16 492 39 16 792 5 164 303 11 474 1 576 961 2 39 199 2357 1081 212 52 9 167 5 56 1 69 30 19 464 17 2844 247 193 890 109 74 3 198 823 21 54 10 64 169 1 0.7776 1.2568 0.0563 0.152 0.3427 0.2804 0.4115 0.3641 0.0345 30.8603 2.9646 2.4587 2.1417 0.1625 0.1412 0.0728 0.508 5e-04 0.0798 99.2725 1.6955 4.9051 1e-04 0.1615 0.3807 0.3063 9.8259 1.9403 1.8686 4.1649 0.3384 2.1183 0.7615 1.5137 0.0023 0.8646 4e-04 1.9353 0.3989 0.1905 0.9861 0.9885 2.7275 0.1701 0.0658 1.0163 0.7579 0.0998 8e-04 3.4558 0.1854 0.5919 0.2499 0.3351 0.0072 5.7304 1e-04 Emergent Canopy Treelet Treelet Treelet Treelet Treelet Canopy Canopy Canopy Emergent Emergent Emergent Treelet Canopy Treelet Treelet Treelet Treelet Emergent Canopy Understory Treelet Treelet Treelet Canopy Emergent Canopy Treelet Understory Understory Canopy Treelet Canopy Canopy Canopy Canopy Canopy Emergent Understory Treelet Canopy Treelet Treelet Treelet Understory Canopy Treelet Treelet Canopy Treelet Canopy Understory Canopy Treelet Canopy Treelet Gui-Cong U&L Gui . L Gui L Gui L Gui-Cong Nig-Cam U&L Gui Africa L Gui L Gui-Cong Gui-Cong L Gui L Gui L Gui-Cong Korup L Gui U&L Gui . L Gui Gui-Cong Africa Africa L Gui L Gui Gui-Cong U&L Gui Gui-Cong L Gui L Gui L Gui Africa L Gui L Gui . Gui-Cong Korup U&L Gui . Gui-Cong L Gui-Cong Africa Gui-Cong Gui-Cong U&L Gui U&L Gui L Gui . Nig-Cam L Gui L Gui-Cong U&L Gui U&L Gui U&L Gui U&L Gui Africa . 123 Biodivers Conserv (2007) 16:2045–2074 2063 Appendix 1 continued Genus Species Family Code N BA Form Range Napoleonaea heudelotii Lecythidaceae NAK5 6 0.0138 Treelet U&L Gui Napoleonaea talbotii Lecythidaceae NATA 135 0.1194 Treelet L Gui Napoleonaea vogelii Lecythidaceae NAPS 26 0.0508 Treelet L Gui Nauclea diderrichii Rubiaceae NADI 19 6.5174 Emergent Gui-Cong Nauclea vanderguchtii Rubiaceae MITR 7 0.0891 Canopy L Gui Newtonia duparquetiana Fabaceae NEDU 78 1.9473 Emergent U&L Gui Newtonia griffoniana Fabaceae NEWI 39 0.0788 Emergent L Gui-Cong Nichallea soyauxii Rubiaceae ROT5 30 0.0079 Treelet Gui-Cong Octoknema affinis Olacaceae OCAF 256 2.4398 Canopy L Gui-Cong Olax latifolia Olacaceae OLAI 71 0.0088 Treelet L Gui Olax triplinerva Olacaceae OLTR 47 0.0061 Treelet L Gui Omphalocarpum elatum Sapotaceae OMPI 16 0.9482 Emergent U&L Gui Opilia sp. Opiliaceae OPIL 4 0.0014 Treelet . Oricia lecomteana Rutaceae ARAL 99 0.0665 Treelet Nig-Cam Oricia renieri Rutaceae RUTA 3 0.0031 Treelet U&L Gui Ormocarpum megalophyllum Fabaceae ORMO 10 0.0016 Treelet U&L Gui Oubanguia alata Lecythidaceae OUAL 14932 218.6065 Canopy L Gui Oubanguia laurifolia Lecythidaceae OULA 93 0.2448 Canopy L Gui Ouratea sp. 1 Ochnaceae OUFB 13 0.0132 Treelet . Oxyanthus formosus Rubiaceae RUAS 2 7e-04 Treelet L Gui Oxyanthus laxiflorus Rubiaceae OXY2 129 0.0575 Treelet L Gui Pachypodanthium sp. Annonaceae PACH 173 0.6091 Emergent . Pachypodanthium staudtii Annonaceae PAST 8 0.6146 Emergent Gui-Cong Panda oleosa Pandaceae PAOL 35 1.59 Canopy L Gui-Cong Pauridiantha afzelii Rubiaceae RUBG 84 0.0083 Treelet U&L Gui Pauridiantha floribunda Rubiaceae PAFL 264 1.5269 Understory L Gui Pauridiantha viridiflora Rubiaceae PSYC 1 0.0043 Treelet L Gui-Cong Pausinystalia macroceras Rubiaceae RUB9 447 5.5646 Canopy L Gui Pavetta rigida Rubiaceae COBN 43 0.0055 Treelet Nig-Cam Pavetta sp. 1 Rubiaceae PANO 74 0.0267 Treelet . Pavetta sp. 2 Rubiaceae RUBN 15 0.0055 Treelet . Pentadesma butyracea Clusiaceae PEBU 66 3.6517 Emergent U&L Gui Pentadesma grandifolia Clusiaceae PENT 30 0.9017 Canopy L Gui Petitiocodon parviflorum Rubiaceae COF2 125 0.0615 Treelet L Gui Phyllanthus sp. Phyllanthaceae RUT2 2 0.0026 Treelet . Phyllanthus sp. Phyllanthaceae RUTS 7 0.1114 Canopy . Phyllobotryon sp. Salicaceae PHYP 8 0.0027 Treelet . Phyllobotryon spathulatum Salicaceae PHSP 26741 12.2868 Treelet L Gui Pierreodendron africanum Simaroubaceae ENT2 7 0.5472 Canopy L Gui Piptadeniastrum africanum Fabaceae PIAF 21 3.4182 Emergent Gui-Cong Piptostigma oyemense Annonaceae PIPI 1032 3.6617 Understory L Gui Placodiscus sp. Sapindaceae PLA 1290 2.0654 Treelet . Pleiocarpa bicarpellata Apocynaceae PLEI 6 0.0052 Treelet L Gui-Cong Pleiocarpa rostrata Apocynaceae PLTA 482 0.1858 Treelet L Gui Poga oleosa Anisophylleaceae POOL 11 13.4212 Emergent L Gui-Cong Polyceratocarpus parviflorus Annonaceae POPA 2048 2.9843 Treelet U&L Gui Polysphaeria macrophylla Rubiaceae POMA 737 0.3906 Treelet U&L Gui Protomegabaria stapfiana Phyllanthaceae PRST 3375 73.2105 Canopy U&L Gui Pseudospondias microcarpa Anacardiaceae PSEU 60 2.3765 Canopy Gui-Cong Psilanthus mannii Rubiaceae RUWS 75 0.0112 Treelet Gui-Cong Psorospermun tenuifolium Clusiaceae PSOR 15 0.0039 Treelet L Gui Psychotria dorotheae Rubiaceae PSYI 925 0.2059 Treelet U&L Gui Psychotria sp. 1 Rubiaceae NPSY 3 9e-04 Treelet . Psychotria sp. 10 Rubiaceae PSYT 1 1e-04 Treelet . Psychotria sp. 2 Rubiaceae PSEL 1 1e-04 Treelet . Psychotria sp. 3 Rubiaceae PSY2 43 0.0067 Treelet . 123 2064 Biodivers Conserv (2007) 16:2045–2074 Appendix 1 continued Genus Species Family Code N BA Form Range Psychotria sp. 4 Rubiaceae PSY3 21 0.0035 Treelet . Psychotria sp. 5 Rubiaceae PSY4 27 0.0056 Treelet . Psychotria sp. 6 Rubiaceae PSY5 6 0.0011 Treelet . Psychotria sp. 7 Rubiaceae PSY6 5 8e-04 Treelet . Psychotria sp. 8 Rubiaceae PSYL 141 0.1145 Treelet . Psychotria sp. 9 Rubiaceae PSYS 43 0.0063 Treelet . Pterocarpus soyauxii Fabaceae DIA4 18 0.0183 Emergent L Gui Ptychopetalum petiolatum Olacaceae PTPE 243 0.9132 Understory L Gui Pycnanthus angolensis Myristicaceae PYAN 25 0.1101 Emergent Africa Pycnocoma macrophylla Euphorbiaceae PYMA 1332 1.0179 Treelet U&L Gui Rauvolfia caffra Apocynaceae RAMA 49 3.786 Canopy Gui-Cong Rauvolfia mannii Apocynaceae VOA2 115 0.0172 Treelet L Gui-Cong Rauvolfia vomitoria Apocynaceae RAVO 56 0.4359 Understory Gui-Cong Rhabdophyllum sp. Ochnaceae RHAB 13 0.0043 Treelet . Rhaptopetalum sp. 1 Lecythidaceae RHA3 47 0.8529 Canopy . Rhaptopetalum sp. 2 Lecythidaceae RHA4 2 7e-04 Treelet . Rhaptopetalum sp. nov. Lecythidaceae RHA2 292 0.7115 Understory Korup Rinorea cf. woermanniana Violaceae RIN9 1049 0.1624 Treelet . Rinorea crassifolia Violaceae RINB 697 0.0909 Treelet Nig-Cam Rinorea dentata Violaceae RIN6 3499 2.6413 Treelet Africa Rinorea gabunensis Violaceae RINC2 4572 1.9577 Treelet L Gui Rinorea kamerunensis Violaceae RINA 3401 0.6102 Treelet L Gui Rinorea leiophylla Violaceae RINI 3404 2.4442 Treelet L Gui Rinorea lepidobotrys Violaceae RILE 5512 9.5489 Treelet U&L Gui Rinorea longicuspis Violaceae NRIN 11 0.0093 Treelet U&L Gui Rinorea oblongifolia Violaceae RIN2 3021 8.3864 Understory Africa Rinorea sp. 2 Violaceae RIN8 193 0.0297 Treelet . Rinorea sp. 3 Violaceae RISP 3 0.0021 Treelet . Rinorea subintegrifolia Violaceae RIN3 4707 1.8189 Treelet Africa Rinorea thomasii Violaceae RINO 461 0.0887 Treelet Nig-Cam Ritchiea sp. Brassicaceae RITC 1 1e-04 Treelet . Rothmannia hispida Rubiaceae ROTH 181 0.3015 Treelet U&L Gui Rothmannia lujae Rubiaceae ROBS 1 0.0119 Understory L Gui-Cong Rothmannia sp. Rubiaceae ROT4 2 5e-04 Treelet . Rothmannia talbotii Rubiaceae ROLU 398 0.2316 Treelet L Gui-Cong Sacoglottis gabonensis Humiriaceae SAGA 2 1.1931 Emergent U&L Gui Salacia lehmbachii Celastraceae SAL2 1106 0.6664 Treelet Gui-Cong Salacia loloensis Celastraceae SALI 739 0.2353 Treelet L Gui Salacia sp. Celastraceae SAAB 1 4e-04 Treelet . Salacia sp. nov. Celastraceae SALS 564 0.085 Treelet L Gui Santiria balsamifera Burseraceae SATR 138 7.304 Emergent U&L Gui Sapium ellipticum Euphorbiaceae EUPI 64 3.0913 Canopy Africa Scaphopetalum blackii Malvaceae SCAI 428 0.0631 Treelet L Gui Schumanniophyton magnificun Rubiaceae SCMA 1425 0.3828 Treelet L Gui-Cong Scottellia klaineana Achariaceae SCCO 599 11.5492 Emergent Gui-Cong Scyphocephalium mannii Myristicaceae SCYP 4 0.0116 Emergent L Gui Scytopetalum klaineanum Lecythidaceae SCKL 200 6.6969 Emergent L Gui Sericanthe auriculata Rubiaceae HEI2 30 0.0341 Treelet L Gui Sibangea similis Putrangivaceae DRSI 713 0.4419 Treelet L Gui Sorindeia grandifolia Anacardiaceae DAMA 12 0.0144 Treelet L Gui Sorindeia juglandifolia Anacardiaceae SOJU 344 1.5937 Understory U&L Gui Soyauxia gabonensis Medusandraceae SOTA 3182 16.7249 Understory L Gui Spathodea campanulata BiGuioniaceae SPCA 1 0.0257 Canopy Africa Staudtia gabunensis Myristicaceae STST 8 1.8503 Emergent L Gui Staudtia kamerunensis Myristicaceae STAU 194 13.4438 Emergent Gui-Cong Sterculia oblonga Malvaceae EROB 5 0.2334 Canopy Gui-Cong 123 Biodivers Conserv (2007) 16:2045–2074 2065 Appendix 1 continued Genus Species Family Code N BA Form Range Sterculia tragacantha Malvaceae STER 38 0.6308 Emergent Gui-Cong Stipularia africana Rubiaceae RUBR 78 0.0111 Treelet U&L Gui Strephonema pseudocola Combretaceae STPO 146 14.217 Emergent U&L Gui Strombosia grandifolia Olacaceae STGR 18 0.4006 Understory U&L Gui Strombosia pustulata Olacaceae STPU 4119 34.4039 Canopy Gui-Cong Strombosia scheffleri Olacaceae STSC 1216 14.0917 Canopy Gui-Cong Strombosia sp. Olacaceae STRO 2523 16.0292 Canopy . Strombosiopsis tetrandra Olacaceae STTE 1763 23.0436 Canopy L Gui-Cong Strychnos sp. Loganiaceae STCO 7 0.6695 Canopy . Symphonia globulifera Clusiaceae SYGL 805 1.463 Emergent Africa Synsepallum stipulatum Sapotaceae SYST 199 2.2383 Understory L Gui-Cong Syzygium guineense Myrtaceae SYGU 5 0.2188 Canopy Africa Syzygium rowlandii Myrtaceae SYRO 72 2.1322 Canopy U&L Gui Tabernaemontana brachyantha Apocynaceae TABR 3583 15.3003 Canopy L Gui Tabernaemontana crassa Apocynaceae TACR 674 2.0902 Canopy Gui-Cong Talbotiella eketensis Fabaceae TAEK 924 10.6727 Canopy Nig-Cam Tapura africana Dichapetalaceae TAAF 418 11.2514 Canopy L Gui Tarenna conferta Rubiaceae RUTL 49 0.0233 Treelet L Gui-Cong Tarenna lasiorachis Rubiaceae RUB8 88 0.0543 Treelet L Gui Tetrapleura tetraptera Fabaceae TETE 8 0.8596 Emergent Gui-Cong Thecacoris leptobotrya Phyllanthaceae EUP2 39 0.0071 Treelet L Gui-Cong Thecacoris sp. Phyllanthaceae THEC 131 1.9599 Canopy . Treculia acuminata Moraceae TREI 133 0.0321 Treelet L Gui Treculia africana Moraceae TRAF 71 0.5562 Emergent Gui-Cong Treculia obovoidea Moraceae TROB 38 1.0111 Canopy L Gui-Cong Tricalysia achoundongiana Rubiaceae RUBP 625 0.1849 Treelet L Gui Tricalysia pangolina Rubiaceae ROT3 13 0.0183 Treelet L Gui Trichilia prieureana Meliaceae TRSP 210 1.0495 Understory Gui-Cong Trichilia sp. Meliaceae TRWE 207 0.5796 Canopy . Trichoscypha acuminata Anacardiaceae TRAC 30 1.725 Canopy L Gui-Cong Trichoscypha klainei Anacardiaceae TRI3 1338 0.6636 Treelet . Trichoscypha patens Anacardiaceae TRI2 1120 1.5909 Understory L Gui Trichoscypha preussii Anacardiaceae TRIC 193 0.619 Understory U&L Gui Trichoscypha sp. Anacardiaceae TRIR 752 0.6728 Treelet . Trichoscypha sp. 1 Anacardiaceae TR22 77 0.0221 Treelet . Trichoscypha sp. 2 Anacardiaceae TRBM 25 1.0108 Emergent . Trichoscypha sp. 3 Anacardiaceae TRHM 147 0.6309 Understory . Trichoscypha sp. 4 Anacardiaceae TRI4 302 0.5262 Understory . Trichoscypha sp. 5 Anacardiaceae TRI72 80 0.2913 Understory . Turraeanthus mannii Meliaceae TUMA 251 0.0563 Treelet Nig-Cam Turreanthus sp. Meliaceae TUMA2 11 0.0017 Treelet . Uapaca acuminata Phyllanthaceae UAAC 6 0.0013 Emergent L Gui Uapaca staudtii Phyllanthaceae UAST 1184 9.6957 Emergent L Gui Urobotrya congolana Opiliaceae EULI 23 0.0103 Treelet Gui-Cong Uvariastrum pynaertii Annonaceae ANNS 446 1.5969 Understory L Gui Uvariodendron sp. Annonaceae UOI 3248 6.5358 Understory . Uvariopsis bakeriana Annonaceae UVBA 1630 0.732 Treelet Nig-Cam Uvariopsis congolana Annonaceae UVCO 4 0.0051 Treelet L Gui Uvariopsis korupensis Annonaceae UVAI 490 1.1131 Understory Nig-Cam Vernonia conferta Asteraceae VECO 9 0.0276 Treelet Gui-Cong Vernonia frondosa Asteraceae VERH 27 0.0611 Treelet U&L Gui Vitex grandifolia Lamiaceae VIGR 181 5.9731 Canopy U&L Gui Vitex sp. 1 Lamiaceae VIT2 77 17.5556 Emergent . Vitex sp. 2 Lamiaceae VIT3 1 1e-04 Emergent . Vitex sp. 3 Lamiaceae VITI 135 7.6492 Emergent . Vitex sp. 5 Lamiaceae VITV 18 0.0134 Treelet . Vitex sp. 6 Lamiaceae VITP 59 0.3192 Canopy . 123 2066 Biodivers Conserv (2007) 16:2045–2074 Appendix 1 continued Genus Species Family Code Voacanga Warneckea Warneckea Warneckea Xylopia Xylopia Xylopia Xylopia Xylopia Zanthoxylum Zanthoxylum Zanthoxylum Zanthoxylum Zeyherella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . psilocalyx jasminoides membranifolia pulcherrima acutiflora aethiopica sp. 1 sp. 2 villosa gilletii sp. 1 sp. 2 sp. 3 letestui . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Apocynaceae Melastomataceae Melastomataceae Melastomataceae Annonaceae Annonaceae Annonaceae Annonaceae Annonaceae Rutaceae Rutaceae Rutaceae Rutaceae Sapotaceae Acanthaceae Sapindaceae Annonaceae Annonaceae Rubiaceae Rubiaceae Rubiaceae Icacinaceae Fabaceae . Rubiaceae Sapindaceae Rubiaceae Fabaceae Rubiaceae Rubiaceae Rubiaceae Ochnaceae Rubiaceae Rubiaceae Rubiaceae Rubiaceae Rubiaceae Rubiaceae Rubiaceae . Rubiaceae Rubiaceae Rubiaceae Chrysobalanaceae . . Sapindaceae VOAI 299 0.0583 Treelet L Gui WAR2 199 0.6043 Understory L Gui-Cong WAME 1857 1.3283 Treelet U&L Gui WARI 240 1.57 Understory L Gui XYAC 245 0.6578 Understory Gui-Cong XYAE 217 8.9498 Canopy Gui-Cong UVAR 55 0.4442 Canopy . XYLO 3 0.1841 Canopy . XYVI 23 0.4367 Canopy U&L Gui ZATE 282 17.7113 Emergent Africa ZANG 3 0.0029 Understory . ZANH 9 0.0023 Treelet . ZATM 19 0.0092 Canopy . ZELE 7 0.1578 Canopy L Gui ACA2 52 0.013 Treelet . ALEL 1 0.5027 Treelet . ANN2 1 0.0015 Treelet . ANNV 2 2e-04 Treelet . COF3 15 0.0025 Treelet . COF4 91 0.1399 Understory . COF5 4 0.0018 Treelet . EUP3 19 0.0068 Treelet . FATD 2 9e-04 Treelet . FLAI 1 6e-04 Treelet . HRUB 1 2e-04 Treelet . LACS 7 0.0012 Treelet . LAS2 1 9e-04 Treelet . LEG2 1 0.0177 Understory . MIT2 3 0.0345 Understory . MOKO 2 0.0685 Understory . NRUB 2 0.0016 Treelet . OURI 184 0.0734 Treelet . RUB3 1 1e-04 Treelet . RUB7 825 0.3924 Treelet . RUBB 109 0.0182 Treelet . RUBM 17 0.3779 Canopy . RUBS 144 0.1162 Treelet . RUBT3 32 0.1122 Treelet . RUBY 7 0.0101 Treelet . RUT3 6 0.0049 Treelet . RUTR 13 0.032 Treelet . RUTR2 4 0.022 Understory . RUYR 2 0.0177 Understory . SOYA 65 0.6595 Canopy . UNID 680 6.4573 . . WDIG 11 0.0036 Treelet . WLOV 1 6e-04 Treelet . N BA Form Range Species names with sp. indicate unnamed but recognizable morphospecies; those with sp. nov. are novel species which have not yet been described. The 4-letter code is our field designation, and is necessary here only for the unidentified species at the end of the table, which otherwise have no designation. All codes but one refer to single morphospecies; the exception, UNID, encompasses all individuals not assigned to one of the morphospecies. N = number of individuals ‡1 cm dbh in 50 ha. BA = basal area in m2 per 50 ha. Range descriptions: Nig-Cam = SE Nigeria and SW Cameroon endemic; L Gui = Lower Guinea endemic; U&L Gui = Upper and Lower Guinea; L Gui-Cong = Lower Guinea-Congolian; Gui-Cong = Guineo-Congolian; Africa = Widespread in Africa. We omit authorities here, but refer instead to http://www.ctfs.si.edu/africatree, where more information is given for all the species 123 Biodivers Conserv (2007) 16:2045–2074 2067 Appendix 2 Genera of the Korup 50-ha plot Genus Family Species N BA Afzelia Aidia Albizia Allexis Allophylus Alsodeiopsis Alstonia Amphimas Angylocalyx Anisophyllea Annickia Anthocleista Anthonotha Antidesma Antrocaryon Aoranthe Aporrhiza Araliopsis Asystasia Aulacocalyx Baikiaea Baillonella Baphia Barteria Beilschmiedia Belonophora Berlinia Bersama Bertiera Blighia Brenania Bridelia Caloncoba Calpocalyx Calycosiphonia Camptostylus Campylospermum Canthium Carapa Carpolobia Casearia Cassipourea Ceiba Cephaelis Chrysobalanus Chrysophyllum Chytranthus Cleistanthus Cleistopholis Coelocaryon Coffea Cola Copaifera Cordia Coula Fabaceae Rubiaceae Fabaceae Violaceae Sapindaceae Icacinaceae Apocynaceae Fabaceae Fabaceae Anisophylleaceae Annonaceae Gentianaceae Fabaceae Phyllanthaceae Anacardiaceae Rubiaceae Sapindaceae Rutaceae Acanthaceae Rubiaceae Fabaceae Sapotaceae Fabaceae Passifloraceae Lauraceae Rubiaceae Fabaceae Melianthaceae Rubiaceae Sapindaceae Rubiaceae Phyllanthaceae Salicaceae Fabaceae Rubiaceae Achariaceae Ochnaceae Rubiaceae Meliaceae Polygalaceae Salicaceae Rhizophoraceae Malvaceae Rubiaceae Chrysobalanaceae Sapotaceae Sapindaceae Phyllanthaceae Annonaceae Myristicaceae Rubiaceae Malvaceae Fabaceae Boraginaceae Olacaceae 2 1 1 1 3 1 1 1 2 3 1 2 4 2 1 1 1 1 1 3 1 1 3 1 7 2 3 1 2 1 1 2 1 1 1 1 4 3 2 1 1 1 1 1 1 2 4 1 2 1 1 23 1 1 1 85 33 1 883 126 16 66 22 5858 222 707 59 419 724 8 79 171 168 8 2735 20 4 2889 79 3070 2548 830 9 725 20 10 59 88 3066 112 143 1032 179 800 485 80 47 2 10 62 193 482 43 49 87 1 70355 8 5 8 5.8927 0.2303 0.509 1.2924 0.0269 0.0022 14.8402 0.8963 3.9654 4.6431 8.3172 3.633 11.9888 2.1656 0.0091 0.5942 0.6366 3.1839 0.0011 3.2458 0.0588 1.0854 17.5033 0.3336 9.5052 0.6971 5.967 0.0048 0.133 0.4155 0.0057 0.8391 6.6273 7.7904 0.1034 0.3376 0.6505 0.316 8.1519 0.1245 0.3792 0.1495 0.0114 0.004 0.4534 0.6871 0.537 0.2203 2.389 5.3102 2e-04 137.8057 0.1553 0.0482 0.3554 123 2068 Biodivers Conserv (2007) 16:2045–2074 Appendix 2 continued Genus Family Species N BA Crateranthus Craterispermum Croton Crotonogyne Crotonogynopsis Crudia Dacryodes Dactyladenia Dasylepis Deinbollia Desbordesia Dialium Dichaetanthera Dichostemma Dicranolepis Didymosalpinx Dinophora Diogoa Diospyros Discoclaoxylon Discoglypremna Dorstenia Dracaena Drypetes Duboscia Endodesmia Englerophytum Engomegoma Entandrophragma Eriocoelum Erismadelphus Erythrina Erythrophleum Erythroxylum Euclinia Eugenia Eurypetalum Fagara Ficus Funtumia Gaertnera Garcinia Gilbertiodendron Glossocalyx Glyphaea Guarea Hallea Heckeldora Heinsia Heisteria Homalium Hoplestigma Hunteria Hylodendron Hymenostegia Lecythidaceae Rubiaceae Euphorbiaceae Euphorbiaceae Euphorbiaceae Fabaceae Burseraceae Chrysobalanaceae Achariaceae Sapindaceae Irvingiaceae Fabaceae Melastomataceae Euphorbiaceae Thymelaeaceae Rubiaceae Melastomataceae Olacaceae Ebenaceae Euphorbiaceae Euphorbiaceae Moraceae Ruscaceae Putrangivaceae Malvaceae Clusiaceae Sapotaceae Olacaceae Meliaceae Sapindaceae Vochysiaceae Fabaceae Fabaceae Rhizophoraceae Rubiaceae Myrtaceae Fabaceae Rutaceae Moraceae Apocynaceae Rubiaceae Clusiaceae Fabaceae Monimiaceae Malvaceae Meliaceae Rubiaceae Meliaceae Rubiaceae Olacaceae Salicaceae Hoplestigmataceae Apocynaceae Fabaceae Fabaceae 1 2 1 1 1 1 2 2 1 3 1 2 1 1 2 1 1 1 14 1 1 1 5 6 1 1 1 1 1 1 1 1 1 1 1 3 1 1 1 1 2 10 1 1 1 3 1 1 1 1 4 1 1 1 3 18 2684 112 3888 2158 1 189 230 2144 616 85 419 3 17255 11 470 2 3220 19143 37 167 10 242 7579 2 270 652 1 44 76 251 8 79 21 131 339 7 109 3 11 721 2828 288 1070 14 149 69 6 103 1174 1287 19 874 13 4279 0.0246 2.7479 3.9922 1.2752 0.9347 0.2489 7.2913 0.7042 7.2305 1.0159 25.0731 5.9826 0.0213 78.804 0.1512 0.5067 2e-04 12.4436 57.2358 0.2272 6.7191 0.001 0.1029 28.2436 0.0012 3.4477 2.4754 0.5555 0.0585 1.5052 13.7765 0.1152 24.5632 1.3316 0.0872 0.1591 0.3198 2.7442 0.0773 0.2239 0.1585 14.3676 2.0869 0.8259 0.0091 7.8527 5.5641 8e-04 0.359 0.1943 15.0165 0.3611 8.462 1.9989 36.5872 123 Biodivers Conserv (2007) 16:2045–2074 2069 Appendix 2 continued Genus Family Species N BA Hypodaphnis Irvingia Isolona Ixora Jollydora Keayodendron Kigelia Klaineanthus Klainedoxa Korupodendron Laccodiscus Lannea Lasianthera Lasianthus Lasiodiscus Lecomtedoxa Lepidobotrys Leptaulus Leptonychia Licania Lophira Macaranga Maesobotrya Maesopsis Magnistipula Mammea Manilkara Maprounea Maranthes Mareya Mareyopsis Margaritaria Massularia Memecylon Microcos Microdesmis Monodora Morelia Morinda Mostuea Musanga Mussaenda Napoleonaea Nauclea Newtonia Nichallea Octoknema Olax Omphalocarpum Opilia Oricia Ormocarpum Oubanguia Ouratea Oxyanthus Lauraceae Irvingiaceae Annonaceae Rubiaceae Connaraceae Euphorbiaceae Bignoniaceae Euphorbiaceae Irvingiaceae Vochysiaceae Sapindaceae Anacardiaceae Icacinaceae Rubiaceae Rhamnaceae Sapotaceae Lepidobotryaceae Icacinaceae Malvaceae Chrysobalanaceae Ochnaceae Euphorbiaceae Phyllanthaceae Rhamnaceae Chrysobalanaceae Clusiaceae Sapotaceae Euphorbiaceae Chrysobalanaceae Euphorbiaceae Euphorbiaceae Phyllanthaceae Rubiaceae Memecylaceae Malvaceae Pandaceae Annonaceae Rubiaceae Rubiaceae Gelsemiaceae Urticaceae Rubiaceae Lecythidaceae Rubiaceae Fabaceae Rubiaceae Olacaceae Olacaceae Sapotaceae Opiliaceae Rutaceae Fabaceae Lecythidaceae Ochnaceae Rubiaceae 1 3 2 2 2 1 1 1 2 1 2 1 1 1 1 1 1 2 2 1 1 1 3 1 3 1 1 1 1 1 1 1 1 6 1 1 1 1 1 1 1 1 3 2 2 1 1 2 1 1 2 1 2 1 2 299 150 342 1998 1168 2 8 2032 15 16 531 16 792 5 164 303 11 475 1537 2 39 199 3650 52 181 56 1 69 30 19 464 17 2844 1516 198 823 21 54 10 64 169 1 167 26 117 30 256 118 16 4 102 10 15025 13 131 16.026 11.4019 1.3131 0.4947 0.6919 0.3641 0.0345 30.8603 5.4233 2.1417 0.3037 0.0728 0.508 5e-04 0.0798 99.2725 1.6955 4.9052 0.5422 0.3063 9.8259 1.9403 6.3719 2.1183 2.2775 0.8646 4e-04 1.9353 0.3989 0.1905 0.9861 0.9885 2.7275 2.1107 3.4558 0.1854 0.5919 0.2499 0.3351 0.0072 5.7304 0.0001 0.184 6.6065 2.0261 0.0079 2.4398 0.0149 0.9482 0.0014 0.0696 0.0016 218.8513 0.0132 0.0582 123 2070 Biodivers Conserv (2007) 16:2045–2074 Appendix 2 continued Genus Family Species N BA Pachypodanthium Panda Pauridiantha Pausinystalia Pavetta Pentadesma Petitiocodon Phyllanthus Phyllobotryon Pierreodendron Piptadeniastrum Piptostigma Placodiscus Pleiocarpa Poga Polyceratocarpus Polysphaeria Protomegabaria Pseudospondias Psilanthus Psorospermun Psychotria Pterocarpus Ptychopetalum Pycnanthus Pycnocoma Rauvolfia Rhabdophyllum Rhaptopetalum Rinorea Ritchiea Rothmannia Sacoglottis Salacia Santiria Sapium Scaphopetalum Schumanniophyton Scottellia Scyphocephalium Scytopetalum Sericanthe Sibangea Sorindeia Soyauxia Spathodea Staudtia Sterculia Stipularia Strephonema Strombosia Strombosiopsis Strychnos Symphonia Synsepallum Annonaceae Pandaceae Rubiaceae Rubiaceae Rubiaceae Clusiaceae Rubiaceae Phyllanthaceae Salicaceae Simaroubaceae Fabaceae Annonaceae Sapindaceae Apocynaceae Anisophylleaceae Annonaceae Rubiaceae Phyllanthaceae Anacardiaceae Rubiaceae Clusiaceae Rubiaceae Fabaceae Olacaceae Myristicaceae Euphorbiaceae Apocynaceae Ochnaceae Lecythidaceae Violaceae Brassicaceae Rubiaceae Humiriaceae Celastraceae Burseraceae Euphorbiaceae Malvaceae Rubiaceae Achariaceae Myristicaceae Lecythidaceae Rubiaceae Putrangivaceae Anacardiaceae Medusandraceae Bignoniaceae Myristicaceae Malvaceae Rubiaceae Combretaceae Olacaceae Olacaceae Loganiaceae Clusiaceae Sapotaceae 2 1 3 1 3 2 1 1 2 1 1 1 1 2 1 1 1 1 1 1 1 11 1 1 1 1 3 1 3 13 1 4 1 4 1 1 1 1 1 1 1 1 1 2 1 1 2 2 1 1 4 1 1 1 1 181 35 349 447 132 96 125 9 26749 7 21 1032 1290 488 11 2048 737 3375 60 75 15 1216 18 243 25 1332 220 13 341 30530 1 582 2 2410 138 64 428 1425 599 4 200 30 713 356 3182 1 202 43 78 146 7876 1763 7 805 199 1.2237 1.59 1.5395 5.5646 0.0377 4.5534 0.0615 0.114 12.2895 0.5472 3.4182 3.6617 2.0654 0.191 13.4212 2.9843 0.3906 73.2105 2.3765 0.0112 0.0039 0.3455 0.0183 0.9132 0.1101 1.0179 4.2391 0.0043 1.5651 27.7907 0.0001 0.5455 1.1931 0.9871 7.304 3.0913 0.0631 0.3828 11.5492 0.0116 6.6969 0.0341 0.4419 1.6081 16.7249 0.0257 15.2941 0.8642 0.0111 14.217 64.9254 23.0436 0.6695 1.463 2.2383 123 Biodivers Conserv (2007) 16:2045–2074 2071 Appendix 2 continued Genus Family Species N BA Syzygium Tabernaemontana Talbotiella Tapura Tarenna Tetrapleura Thecacoris Treculia Tricalysia Trichilia Trichoscypha Turraeanthus Turreanthus Uapaca Urobotrya Uvariastrum Uvariodendron Uvariopsis Vernonia Vitex Voacanga Warneckea Xylopia Zanthoxylum Zeyherella Myrtaceae Apocynaceae Fabaceae Dichapetalaceae Rubiaceae Fabaceae Phyllanthaceae Moraceae Rubiaceae Meliaceae Anacardiaceae Meliaceae Meliaceae Phyllanthaceae Opiliaceae Annonaceae Annonaceae Annonaceae Asteraceae Lamiaceae Apocynaceae Melastomataceae Annonaceae Rutaceae Sapotaceae 2 2 1 1 2 1 2 3 2 2 10 1 1 2 1 1 1 3 2 6 1 3 5 4 1 77 4257 924 418 137 8 170 242 638 417 4064 251 11 1190 23 446 3248 2124 36 471 299 2296 543 313 7 2.351 17.3905 10.6727 11.2514 0.0776 0.8596 1.967 1.5994 0.2032 1.6291 7.7526 0.0563 0.0017 9.697 0.0103 1.5969 6.5358 1.8502 0.0887 31.5106 0.0583 3.5026 10.6726 17.7257 0.1578 N = number of individuals ‡1 cm dbh in 50 ha. BA = basal area in m2 per 50 ha Appendix 3 Families of the Korup 50-ha plot Family Genera Species N BA Acanthaceae Achariaceae Anacardiaceae Anisophylleaceae Annonaceae Apocynaceae Asteraceae Bignoniaceae Boraginaceae Brassicaceae Burseraceae Celastraceae Chrysobalanaceae Clusiaceae Combretaceae Connaraceae Dichapetalaceae Ebenaceae Euphorbiaceae Fabaceae Gelsemiaceae Gentianaceae 1 3 5 2 11 7 1 2 1 1 2 1 5 6 1 1 1 1 14 24 1 1 2 3 15 4 22 11 2 2 1 1 3 4 9 16 1 2 1 14 14 39 1 2 60 2886 4504 233 10744 6215 36 9 5 1 327 2410 570 4070 146 1168 418 19143 27798 19393 64 59 0.0141 19.1173 11.8191 18.0643 41.1381 45.405 0.0887 0.0602 0.0482 0.0001 14.5953 0.9871 4.7998 24.7002 14.2170 0.6919 11.2514 57.2358 132.3382 143.6448 0.0072 3.6330 123 2072 Biodivers Conserv (2007) 16:2045–2074 Appendix 3 continued Family Genera Species N BA Hoplestigmataceae Humiriaceae Icacinaceae Irvingiaceae Lamiaceae Lauraceae Lecythidaceae Lepidobotryaceae Loganiaceae Malvaceae Medusandraceae Melastomataceae Meliaceae Melianthaceae Memecylaceae Monimiaceae Moraceae Myristicaceae Myrtaceae Ochnaceae Olacaceae Opiliaceae Pandaceae Passifloraceae Phyllanthaceae Polygalaceae Putrangivaceae Rhamnaceae Rhizophoraceae Rubiaceae Ruscaceae Rutaceae Salicaceae Sapindaceae Sapotaceae Simaroubaceae Thymelaeaceae Urticaceae Violaceae Vochysiaceae 1 1 3 3 1 2 5 1 1 8 1 3 7 1 1 1 3 4 2 4 9 2 2 1 9 1 2 2 2 38 1 4 4 8 8 1 1 1 2 2 1 1 5 6 6 8 10 1 1 32 1 5 11 1 6 1 5 5 5 8 13 2 2 1 16 1 7 2 2 86 5 8 9 19 9 1 2 1 14 2 19 2 1302 250 471 3369 15751 11 7 72579 3182 2301 1678 9 1516 1070 255 318 416 1281 14659 27 858 79 9237 485 8292 216 68 22953 242 692 28204 3321 1375 7 11 169 31413 267 0.3611 1.1931 5.4222 41.8983 31.5106 25.5312 227.3219 1.6955 0.6695 142.7527 16.7249 3.5241 17.7510 0.0048 2.1107 0.8259 1.6777 20.726 2.5101 10.5673 104.8857 0.0117 1.7754 0.3336 95.5739 0.1245 28.6855 2.1981 1.4811 35.7871 0.1029 23.7234 34.3125 7.0107 106.8651 0.5472 0.1512 5.7304 29.0831 15.9182 N = number of individuals ‡1 cm dbh in 50 ha. BA = basal area in m2 per 50 ha References Angiosperm Phylogeny Group (2003) An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II. Bot J Linn Soc 141:399–436 Aubréville A, Leroy J-F, Morat P (eds) (1961–1991) Flore du Gabon, vols 1–32. Muséum National d’Histoire Naturelle, Paris Aubréville A, Leroy J-F, Morat P, Satabie B, Achoundong G (eds) (1963–2001) Flore du Cameroun, vols 1–37. Muséum National d’ Histoire Naturelle, Paris and Ministère de la Récherche Scientifique et Technique, Yaoundé, Cameroun Burgess N, Küper W, Mutke J, Brown J, Westaway S, Turpie S, Meshack C, Tapling J, McClean C, Lovett JC (2005) Major gaps in the distribution of protected areas for threatened and narrow range Afrotropical plants. Biodivers Conserv 14:1877–1894 Brown JH (1995) Macroecology. University of Chicago Press, Chicago, USA 123 Biodivers Conserv (2007) 16:2045–2074 2073 Chuyong GB, Condit R, Kenfack D, Losos E, Sainge SM, Songwe NC, Thomas DW (2004) Korup Forest Dynamics Plot, Cameroon. In: Losos EC, Leigh Jr EG (eds) Forest diversity and dynamism: findings from a large-scale plot network. University of Chicago Press, Chicago, pp 506–516 Condit R (1998) Tropical forest census plots. Springer-Verlag, NY Condit R, Hubbell SP, Foster RB (1996a) Changes in a tropical forest with a shifting climate: results from a 50 ha permanent census plot in Panama. J Trop Ecol 12:231–256 Condit R, Hubbell SP, LaFrankie JV, Sukumar R, Manokaran N, Foster RB, Ashton PS (1996b) Species-area and species-individual relationships for tropical trees: a comparison of three 50 ha plots. J Ecol 84:549–562 Condit R, Robinson WD, Ibáñez R, Aguilar S, Sanjur A, Martı́nez R, Stallard R, Garcı́a T, Angehr G, Petit L, Wright SJ, Robinson TR, Heckadon S (2001) Maintaining the canal while conserving biodiversity around it: a challenge for economic development in Panama in the 21st century. Bioscience 51:135–144 Condit R, Ashton P, Baslev H, Brokaw N, Bunyavejchewin S, Chuyong G, Co L, Dattaraja HS, Davies S, Esufali S, Ewango CEN, Foster R, Gunatileke N, Gunatileke S, Hernandez C, Hubbell S, John R, Kenfack D, Kirakiprayoon S, Hall P, Hart T, Itoh A, Lafrankie J, Liengola I, Lagunzad D, Lao S, Losos E, Magard E, Makana J, Manokaran N, Navarette H, Mohammed Nur S, Okhubo T, Pérez R, Smaper C, Hua Seng L, Sukumar R, Svenning JC, Tan S, Thomas D, Thomson J, Valleho M, Villa Muñoz G, Valencia R, Yamakura T, Zimmerman J (2005) Tropical tree alpha-diversity: results from a worldwide network of large plots. Biol Skr 55:565–582 Connell JH, Lowman MD (1989) Low-diversity tropical rain forests: some possible mechanisms for their existence. Am Nat 134:88–119 Gartlan JS, Newbery DM, Thomas DW, Waterman PG (1986) The influence of topography and soil phosphorus on the vegetation of Korup Forest Reserve, Cameroon. Vegetatio 65:131–148 Gaston KJ (2003) The structure and dynamics of geographic ranges. Oxford University Press, Oxford, UK Gereau ER, Kenfack D (2000) Uvariopsis korupensis, Annonaceae nouvelle du Cameroun. Adansonia 22:39–43 Hart T (1990) Monospecific dominance in tropical rain forests. Trends Ecol Evol 5:6–10 Hutchinson J, Dalziel JM, Hepper FN (1954–1972) Flora of west tropical Africa, vols. 1–3, 2nd edn. Crown Agents, London Kenfack D, Ewango C, Thomas DW (2004) Manilkara lososiana, a new species of Sapotaceae from Cameroon. Kew Bull 59:609–612 Kenfack D, Sainge NM, Thomas DW (2006) A new species of Cassipourea (Rhizophoraceae) from western Cameroon. Novon 16:62–65 Küper W, Sommer JH, Lovett JC, Mutke J, Linder HP, Beentje HJ, Van Rompaey R, Chatelain C, Sosef M, Barthlott W (2004) Africa’s hotspots of biodiversity redefined. Ann Miss Bot Garden 91(4):525–535 Letouzey R (1968) Étude Phytogeogéographique du Cameroun. Encyclopédie Biologique 69. Éditions Paul Lechevalier, Paris Letouzey R (1985) Notice de la carte phytogéographique du Cameroun au 1:500 000. Inst Carte Int Végét (Toulouse, France) 4:95–142 Linder PH (2001) Plant diversity and endemism in sub-Saharan tropical Africa. J Biogeogr 28:160–182 Lovett JC, Rudd S, Taplin J, Frimodt-Moller C (2000) Patterns of plant diversity in Africa south of the Sahara and their implications for conservation management. Biodivers Conserv 9:37–46 Makana J-R, Hart T B, Hart JA (1998) Forest structure and diversity of lianas and understory treelets in monodominant and mixed stands in the Ituri Forest, Democratic Republic of the Congo. In: Dallmeier F, Comiskey JA (eds) Forest biodiversity diversity research, monitoring, and modeling. UNESCO, The Parthenon Publishing Group, Paris Makana J-R, Hart TB, Liengola I, Ewango C, Hart JA, Condit R (2004) Ituri forest dynamics plot, Democratic Republic of Congo. In: Losos EC, Leigh Jr EG (eds) Forest diversity and dynamism: findings from a large-scale plot network. University of Chicago Press, Chicago, pp 492–505 Maley J (1987) Fragmentation de la forêt dense humide africaine et extension des biotopes montagnards au quaternaire récent: nouvelles données polliniques et chronologiques. Implications paleoclimatiques et biogeographiques. Paleoecol Afr 18:307–334 Manokaran N, LaFrankie JV, Kochummen KM, Quah ES, Klahn J, Ashton PS, Hubbell SP (1992) Stand table and distribution of species in the 50-ha research plot at Pasoh Forest Reserve. Forest Research Institute of Malaysia, Kepong, Malaysia 123 2074 Biodivers Conserv (2007) 16:2045–2074 Marimon BS, Felfili MJ, Haridasan M (2001) Studies in monodominant forests in eastern Mato Mrosso, brazil: i. A forest of Brosimum rubescens Taub. Edin J Bot 58:123–137 Nascimento MT, Proctor J (1997) Population dynamics of five tree species in a monodominant Peltogyne forest and two other forest types on Maraca Island, Roraima, Brazil. For Ecol Manage 94:15–128 Newbery DM, Gartlan JS (1996) A structural analysis of rain forest at Korup and Douala-Edea, Cameroon. Proc R Soc Edin 104B:177–224 Newbery DM, van der Burgt XM, Moravie M-A (2004) Structure and inferred dynamics of a large grove of Microberlinia bisulcata trees in central African rain forest: the possible role of periods of multiple disturbance events. J Trop Ecol 20:131–143 Pitman NCA, Terborgh J, Silman MR, Muñez PV (1999) Tree species distributions in an upper Amazonian forest. Ecology 80:2651–2661 Richards PW (1973) Africa: the odd man out. In: Meggers BJ, Ayensu ES, Duckorth WD (eds) Tropical forest ecosystems in Africa and South America: a comparative review. Smithsonian Institution Press, Washington D.C., pp 21–26 Rodrigues ASL, Gaston K (2000) Rarity and conservation planning across geopolitical units. Conserv Biol 16:674–682 Rodrigues ASL, Andelman SJ, Bakarr MI, Boitani L, Brooks TM, Cowling RM, Fishpool LDC, da Fonseca GAB, Gaston KJ, Hoffmann M, Long JS, Marquet PA, Pilgrim JD, Pressey RL, Schipper J, Sechrest W, Stuart SN, Underhill LG, Waller RW, Watts MEJ, Yan X (2004) Effectiveness of the global protected area network in representing species diversity. Nature 428: 640–643 Ruokolainen K, Tuomisto H, Chave J, Muller-Landau HC, Condit R, Pitman N, Terborgh J, Hubbell SP, Leigh Jr EG, Duivenvoorden JF, Svenning J-C, Wright SJ (2002) Beta-diversity in tropical forests. Science 297:1439 Sankovski A, Pridnia M (1995) A comparison of the Southern Appalachian (U.S.A.) and Southwestern Caucasus (Russia) forests: influences of historical events and present environment. J Biogeogr 22:1073–1081 Shaw DC, Franklin JF, Bible K, Klopatek J, Freeman E, Greene S, Parker GG (2004) Ecological setting of the Wind River old-growth forest. Ecosystems 7:427–439 Sonké B, Kenfack D, Robbrecht E (2002) A new species of the Tricalysia atherura group (Rubiaceae) from southwestern Cameroon. Adansonia 24:173–177 Thomas DW, Kenfack D, Chuyong GB, Moses SN, Losos E, Condit RS, Songwe NC (2003) Tree species of southwestern Cameroon: tree distribution maps, diameter tables, and species documentation of the 50-hectare Korup Forest dynamics plot. Center for Tropical Forest Science of the Smithsonian Tropical Research Institute, Washington D.C. Valencia R, Foster RB, Villa G, Condit R, Svenning JC, Hernández C, Romoleroux K, Losos E, Magård E, Balslev H (2004) Tree species distributions and local habitat variation in the Amazon: a large forest plot in eastern Ecuador. J Ecol 92:214–229 White F (1979) The Guineo-Congolian Region and its relationships to other phytochoria. Bull Jardin Bot Natl Belg 49:11–55 White F (1983) The vegetation of Africa. UNESCO, Paris, 356 pp 123

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