contribuţii botanice 2014, xlix: 143-178 grădina botanică...

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Contribuţii Botanice 2014, XLIX: 143-178 Grădina Botanică “Alexandru Borza” Cluj-Napoca FOREST TYPES AND BIODIVERSITY AROUND THE GREAT RIFT VALLEY IN KENYA Kazue FUJIWARA 1 , Takuya FURUKAWA 2 , Samuel KIBOI 3 , Simon MATHENGE 3 , Patrick MUTISO 3 , Hisanori HAYASHI 4 , Shin-ichi MEGURO 4 1 Yokohama City University, Graduate School of Nanobioscience, Seto 22-2, Kanazawa-ku, Yokohama, 236-0027 Japan 2 Forestry and Forest Products Research Institute, University of Nairobi, Kenya 3 University of Nairobi, Kenya 4 Global Environmental Strategies, Japanese Center for International Studies in Ecology (IGES-JISE) e-mail: [email protected] Abstract: The forests along the Great Rift Valley in Kenya are unique as green islands in dry land and extend from southern Ethiopia to Tanzania. These forests include Afromontane and dry-tropical forests, studied in Kenya by Bussmann and Beck, and in southern Ethiopia by Bussmann. Two taxonomical classes (Ocotetea usambarensis in moist climate and Juniperetea procerae at a dry site) were described prior to our phytosociological studies, which were carried out on the east and west sides of the Great Rift Valley in 2006-2014, and results were compared with Bussmann's outcome from the 1990s. New results are presented in this paper: 1) Forests remain on the west side above 2000 m and are characterized by Ocotea kenyensis (Lauraceae); these are different from the moist Afromontane forest at an eastern site (Ocotetea usambarensis) next to the Rift Valley. Four months with > 100 mm rainfall occur from March or April to August or September. 2) At 1300-2200 m at the eastern site, there are unique dry forests characterized by Brachylaena huilliensis (Compositae), Warburgia ugandensis (Canellaceae), Elaeodendron buchananii (Celastraceae), Vepris trichocarpa (Rutaceae), Calodendrum capense, etc. Some of these are unique species and extend to the rainforest in South Africa. 3) Higher-altitude forests at the eastern site are characterized by Podocarpus latifolius and Juniperus procera, which belong to the Juniperetea procerae. Podocarpus latifolius forests occur at the western site, but these are different from those at the eastern site. 4) The forests of the isolated northern mountains Marsabit and Kulal are different; there is no Podocarpus or Juniperus, and higher moisture permits hanging Usnea usneoides to occur at Marsabit. 5) The forest area of Kenya was potentially 12%, and remnant natural forests only cover 1.9%. This small area permits various forest types along a moisture gradient. 6) Several woody and fern genera of the Afromontane forests are common to tropical montane and subtropical forests in Asia, including Podocarpus, Juniperus, Myrsine, Rapanea, Psychotria, Rumohra, Asplenium, Pteris, and Dryopteris. 7) Our and Bussmann's classifications are not the same at the association or community level. There is thus the question whether the 1990s data were from a quite different area or whether some vegetation disappeared due to human or wildlife affects. Keywords: Afromontane forests, dry montane forests, new syntaxa, phytosociological study, tropical dry forests. Plant names: Turill et al. (1952-2006); Knox et al. (1994). Introduction Phytosociological study of forests in Kenya was carried out in great detail by Bussmann and Beck [5] and by Bussmann [4]. The present authors took new vegetation data in 2006-2014, for understanding Kenyan forests, for restoration of natural forests in degraded areas, and for scientific interest in similar genera occurring in subtropical laurel forests in Asia and Afromontane forests in

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  • Contribuţii Botanice – 2014, XLIX: 143-178

    Grădina Botanică “Alexandru Borza”

    Cluj-Napoca

    FOREST TYPES AND BIODIVERSITY AROUND

    THE GREAT RIFT VALLEY IN KENYA

    Kazue FUJIWARA1, Takuya FURUKAWA

    2, Samuel KIBOI

    3, Simon MATHENGE

    3, Patrick MUTISO

    3,

    Hisanori HAYASHI4, Shin-ichi MEGURO

    4

    1 Yokohama City University, Graduate School of Nanobioscience, Seto 22-2, Kanazawa-ku,

    Yokohama, 236-0027 Japan 2 Forestry and Forest Products Research Institute, University of Nairobi, Kenya

    3 University of Nairobi, Kenya

    4 Global Environmental Strategies, Japanese Center for International Studies in Ecology (IGES-JISE)

    e-mail: [email protected]

    Abstract: The forests along the Great Rift Valley in Kenya are unique as green islands in dry land and extend

    from southern Ethiopia to Tanzania. These forests include Afromontane and dry-tropical forests, studied in Kenya by

    Bussmann and Beck, and in southern Ethiopia by Bussmann. Two taxonomical classes (Ocotetea usambarensis in moist

    climate and Juniperetea procerae at a dry site) were described prior to our phytosociological studies, which were carried

    out on the east and west sides of the Great Rift Valley in 2006-2014, and results were compared with Bussmann's

    outcome from the 1990’s. New results are presented in this paper: 1) Forests remain on the west side above 2000 m and

    are characterized by Ocotea kenyensis (Lauraceae); these are different from the moist Afromontane forest at an eastern

    site (Ocotetea usambarensis) next to the Rift Valley. Four months with > 100 mm rainfall occur from March or April to

    August or September. 2) At 1300-2200 m at the eastern site, there are unique dry forests characterized by Brachylaena

    huilliensis (Compositae), Warburgia ugandensis (Canellaceae), Elaeodendron buchananii (Celastraceae), Vepris

    trichocarpa (Rutaceae), Calodendrum capense, etc. Some of these are unique species and extend to the rainforest in

    South Africa. 3) Higher-altitude forests at the eastern site are characterized by Podocarpus latifolius and Juniperus

    procera, which belong to the Juniperetea procerae. Podocarpus latifolius forests occur at the western site, but these are

    different from those at the eastern site. 4) The forests of the isolated northern mountains Marsabit and Kulal are

    different; there is no Podocarpus or Juniperus, and higher moisture permits hanging Usnea usneoides to occur at

    Marsabit. 5) The forest area of Kenya was potentially 12%, and remnant natural forests only cover 1.9%. This small

    area permits various forest types along a moisture gradient. 6) Several woody and fern genera of the Afromontane

    forests are common to tropical montane and subtropical forests in Asia, including Podocarpus, Juniperus, Myrsine,

    Rapanea, Psychotria, Rumohra, Asplenium, Pteris, and Dryopteris. 7) Our and Bussmann's classifications are not the

    same at the association or community level. There is thus the question whether the 1990s data were from a quite

    different area or whether some vegetation disappeared due to human or wildlife affects.

    Keywords: Afromontane forests, dry montane forests, new syntaxa, phytosociological study, tropical dry

    forests.

    Plant names: Turill et al. (1952-2006); Knox et al. (1994).

    Introduction

    Phytosociological study of forests in Kenya was carried out in great detail by Bussmann and

    Beck [5] and by Bussmann [4]. The present authors took new vegetation data in 2006-2014, for

    understanding Kenyan forests, for restoration of natural forests in degraded areas, and for scientific

    interest in similar genera occurring in subtropical laurel forests in Asia and Afromontane forests in

  • 144 K. FUJIWARA, T. FURUKAWA, S. KIBOI, S. MATHENGE, P. MUTISO,

    H. HAYASHI, SHIN-ICHI MEGURO

    eastern Africa. Africa separated from Gondwana as the supercontinent split up in the middle

    Mesozoic era. The Great Rift Valley resulted from this tectonic separation, accompanied by

    volcanic activity [36, 43], Nowadays, remaining natural forests cover only 1.9% of the total area of

    Kenya [40]. With such small areas of remnant natural forests, it is difficult to find good natural

    forests and to describe their species composition for the future. We looked for natural forests with

    less human impact (e.g. livestock) and less disturbance by wildlife, which overpopulates some

    forest areas. Forests in Kenya were described by White [42] and Bussman & Beck [5] as: 1)

    Savanna with Acacia on lower and dry sites; 2) tropical dry forests with evergreen and semi-

    evergreen bushland and thicket on the middle slopes; 3) Afromontane moist forest and dry conifer

    forests; 4) Hagenia forest at timberline; and 5) ericaceous scrub above timberline. In this paper we

    tried: 1) to clarify the forest types of Kenya and compare with data from Bussmann and Beck [5]

    and Bussmann [4]; 2) to understand characteristics of tropical dry forests and Afromontane forests,

    especially evergreen and semi-evergreen forests; and 3) to understand differences of Kenyan forest

    types on the east and west sides of the Great Rift Valley.

    Material and Method

    Study area

    The study area is from 1230 m to 3405 m altitude, from 2º40’00.1” N to 1º53’48.8” S

    latitude and from 38º00’01.11” to 35º52’01.2” E longitude, along the Great Rift Valley of western

    Kenya (Fig. 1, Table 1 in Appendix). We used climate data from the world climate database of E.O.

    Box (University of Georgia, USA). The climates of the study sites are dry tropical and moist

    mountain climates, with 604-1165 mm precipitation except at the mountain tops, where

    precipitation reaches 1200-2000 mm (see Figure 1; [4], cf. [9]); there are one or two drier seasons

    per year. The mean temperature is 16 to 23ºC and the temperature of the coldest month is

    sometimes below freezing at 2800 m in Mau. A drier climate occurs in the eastern lower hills of the

    Great Rift Valley.

    The topography of the Great Rift Valley area involves continuous hilly area at the western

    site and solitary mountains such as Mt. Kenya, Mt. Marsabit and Mt. Kural, plus hills of the

    Mattews Range, the Karrissa and Loita Hills, Karura and Ngong. Various rock types underlie these

    areas, but granite and volcanic rocks are the main substrates. The main soil types are Vertisols,

    Regosols, Lithosols and Cambisols on the plains, with humic Nitisols and Acrisols on the base

    substrates, volcanic areas on the mountain slopes, and deep humic Andosols above about 2700 m

    [4, 30, 31, 32, 37].

    Vegetation survey

    The vegetation survey was carried out in 2006-2014, based on Braun-Blanquet and Fujiwara

    [1, 10]. Forest description involved: 1) selecting homogeneous sites with less human impact, as by

    livestock or illegal cutting; and, 2) describing the species composition in four or five forest layers

    (estimating by height and density) and assigning cover and occurrence frequency. Environmental

    variables recorded include elevation, slope, aspect and inclination, etc. (Table 2 in Appendix) and

    vegetation profiles.

    Data analysis

    The classification of plant communities was carried out by Braun-Blanquet methodology.

    Data from Bussmann and Beck [5] and Bussmann [4] were added into the synoptic table

  • FOREST TYPES AND BIODIVERSITY AROUND THE GREAT RIFT VALLEY… 145

    and compared, but environmental data are missing for the Bussmann sites.

    De-trended Correspondence Analysis (DCCA, PC-CORD version 4.25) was used to analyze

    the relationship between plant communities and environmental variables, using the middle values of

    cover-abundance on the Braun-Blanquet scale. Elevation, slope, minimum temperature of coldest

    months, precipitation, latitude, longitude, and times of drier periods in a year, from the authors'

    relevé data, were selected as the main variables to represent environmental conditions.

    Fig. 1: Average annual rainfall and temperature in Kenya.

    Results

    The results consist mainly of a phytosociological summary. Here we discuss the

    characteristic distribution of tropical dry forests and Afromontane forests. Therefore a new

    syntaxonomical classification is attempted, as a basis for discussion.

    I. Tropical dry forests

    Tropical dry forests in Kenya were described by several authors [41, 23, 5, 4, 16]. The main

    canopy species of Kenyan tropical dry forests are Croton megalocarpus, Celtis africana,

    Fagaropsis angolensis, Calodendrum capense, Grewia similis, and Euclea divinorum. These

  • 146 K. FUJIWARA, T. FURUKAWA, S. KIBOI, S. MATHENGE, P. MUTISO,

    H. HAYASHI, SHIN-ICHI MEGURO

    species are common from Mt. Marsabit and Mt. Kulal (northern Kenya) to the Loita Hills (southern

    Kenya). The main forests are characterized by Elaeodendron buchananii, Vepris trichocarpa,

    Brachylaena huillensis, Warburgia ugandensis, Rawsonia lucida, and Manilkara discolor,

    especially in the Nairobi area.

    These tropical dry forests were assigned to Juniperetea procerae Bussmann et Beck 1995,

    but there are no common characteristic species in Juniperus procera and tropical dry forests (Table

    3 in Appendix). In this paper, we propose a new class for Kenyan tropical dry forests.

    1. Schoenoxiphio lehmannii-Crotonietea megalocarpae cls. nova (Table 3 in Appendix)

    1-1. Schoenoxiphio lehmannii-Crotonietalia megalocarpae ord. novus (Table 3 in Appendix)

    Diagnostic taxa: Strychnos usambarensis, Barleria micrantha, Schoenoxiphium lehmannii, Setaria

    plicatilis, Ehretia cymosa, Phaulopsis imbricata, Croton megalocarpus, Clausena anisata, Celtis

    africana, Fagaropsis angolensis, Ficus thonningii, Commelina benghalensis, Allophylus ferruginus,

    Calodendrum capense, Grewia similis, Acokanthera schimperi, Euclea divinorum, Zanthoxylum

    usambarense, Ehrharta erecta var. abyssinica, Ochna ovata, Maytenus undata, Vangueria infausta.

    Holotype: Brachylaenion huillensis Bussmann et Beck 1995.

    Forests dominated by Croton megalocarpus are mostly secondary forests but occur also as a

    zone next to dry savanna areas. Tropical dry forests contain C. megalocarpus relatively frequently,

    except at Mt. Kulal. The canopy trees are usually very tall, at 20-40 m; only forests in the Loita

    Hills and Karrisa Hills are 12-20 m. The canopy is relatively open, with most forests having 50-

    70% canopy cover but some forests reaching 80%. The occurrence of C. megalocarpus in open

    habitats matches its character as a light-demanding, fast-growing tree [22]. A similar character

    species is Brachylaena huillensis, whose seedlings demand light but do grow especially under

    mother trees. Seed viability decreases under sunlight exposure, but continuous regeneration was

    found in both disturbed forest stands and under closed canopies [24].

    1-1-1. Strychnoso mitis-Diospyrosion abyssinicae all. nova (Table 3 and 4 in Appendix)

    Diagnostic taxa: Strychnos mitis, Dorstenia brownia, Rinorea convallarioides ssp. marsabitensis,

    Dregea abyssinica.

    Holotype: Pavetto gardeniifoliae-Cassipouretum malosanae ass. nova

    Mt. Marsabit, Mt. Kulal and Kitichii Camp in the Mattews Range are ‘isolated islands' in the

    savanna. These are volcanic mountains, with more humid environments. In particular, Usnea

    usnoides is hanging on the trees in the Marsabit forests surrounding the Paradise Lake at the top of

    the National Park. These are classified as a new alliance with the diagnostic taxa listed above. Three

    new associations are contained.

    1-1-1-1. Pavetto gardeniifoliae-Cassipouretum malosanae ass. nova (Table 4 in Appendix)

    Diagnostic taxa: Pavetta gardeniifolia, Isoglossa laxa, Scadoxus multiflorus ssp. multiflorus,

    Oxyanthus speciosus ssp. stenocarpus, Margaritaria discoidea, Allophylus abyssinica.

    Holotype: running no. 3 (Kl-3: 1946 m).

    Mt. Kulal is an isolated volcanic mountain. Lake Turkana lies on its west side, and the

    surrounding area is covered by volcanic gravel. A wide area of the forest is protected on top of the

    mountain. The height of this forest is 27-40 m, and canopy trees have diameters (DBH) of 20-47

    cm. Five or six species are mixed in the tall canopy layer. Four to five layers are recognized,

  • FOREST TYPES AND BIODIVERSITY AROUND THE GREAT RIFT VALLEY… 147

    according to presence of very tall canopy trees such as Cassipourea molasana, Olea europea ssp.

    africana, Strychnos usambariensis, Diospyros abyssinica, Ocotea kenyensis and Ficus thonningii.

    The forest of Mt. Kulal is protected by the local Conservation Committee, but cattle can be seen

    grazing everywhere. Also, baboons disturb the soil to search for soil roaches. Relevé data were

    recorded at 1835-1946 m elevation. Ocotea kenyensis occurs on Mt. Kulal, but the other diagnostic

    taxa (e.g. Tabernaemontana, Sysygium, Psychotria, and Piper) are missing. Croton megalocarpus is

    also missing on top of the mountain.

    1-1-1-2. Veprisio hanagensi-Drypetetum gerrardii ass. nova (Table 4 in Appendix)

    Diagnostic taxa: Vepris hanagensis, Erythroxylum fischeri, Ritchiea albersii, Coffea arabica,

    Premna maxima.

    Holotype: running no. 12 (Mar-6: 1392 m).

    The forests in Marsabit National Park are relatively well preserved at 1230-1540 m. The

    striking characteristic species is Usnea usneoides hanging on canopy trees surrounding Lake

    Paradise. The height of the forests is 27-38 m, and average canopy cover is about 63% (50-80%). It

    is a relatively open forest. The forests in Marsabit are characterized by fewer species than other

    forests, and cover by Drypetes gerardii is relatively high (20-50%). Githae (2006) made an

    inventory and diversity assessment in the forest dominated by Drypetes gerardii. Mt. Marsabit is

    missing some species from tropical dry forests, such as Celtis africana and Vepris simplicifolia. The

    mean number of species is 30 (21-41), which is relatively low. Bussmann [4] reported the presence

    of Coffeo arabicae-Rinoreetum convallarioidis, but its diagnostic taxa were not recognized in our

    vegetation sampling; also, tropical dry-forest species are missing in his sampling (Table 3: running

    no. 31).

    1-1-1-3. Argomuellero macrophylli-Crotonetum megalocarpae ass. nova (Table 4 in Appendix)

    Diagnostic taxa: Allophylus rubifolius, Erythroxylum emarginatum, Argomuellera macrophylla,

    Vangueria madagascariensis, Haplocoelum foliolosum, Encephalartos tegulaneus,

    Pseuderanthemum hildebrandtii, Sapium ellipticum, Bauhinia tomentosa, Craibia laurentii,

    Hippocratea paniculata

    Holotype: Ken-90 (running no. 25)

    The forest height is lower, at 25-33 m, than at Mt. Marsabit and Mt. Kulal; canopy cover is

    50-80%. Characteristic is the dominance of Aegollera macrophylla (Euphorbiaceae) in the shrub

    layer and of Encephalartos tegulaneus, the Kenyan giant cycad (Zamiaceae), in the shrub layer to

    sub-canopy tree layer. The physiognomy of the forest is different from that of other tropical dry

    forests.

    1-1-2. Brachylaenion huillensis Bussmann et Beck 1995 (Table 3 in Appendix)

    Diagnostic taxa: Elaeodendron buchananii, Vepris trichocarpa, Brachylaena huillensis, Asparagus

    setaceus, Pterolobium stellatum, Vernonia holstii, Warburgia ugandensis, Dombeya burgessiae,

    Strychnos henningsii, Rawsonia lucida, Manilkara discolor, Uvaria scheffleri, Chrysophyllum

    viridifolium

    Holotype: Elaeodendro buchananii-Brachylaenetum huillensis Bussman et Beck 1995 (Table 3 in

    Appendix)

  • 148 K. FUJIWARA, T. FURUKAWA, S. KIBOI, S. MATHENGE, P. MUTISO,

    H. HAYASHI, SHIN-ICHI MEGURO

    This alliance is typical of tropical dry forests in Kenya. The diagnostic taxa occur in

    southern Kenya (Karura, Ngong, Oloolua in Nairobi, and the Loita Hills) and on Mt. Kenya [5].

    Most forests are 19-25 m tall, except at Karura, which still has well preserved forests 23-40 m tall.

    Forests in the Loita Hills are shorter than other forests, at 12-19 m (an exception is 27 m). Two new

    suballiances are distinguished but still need more discussion.

    1-1-2-1. Crotonenion alieno-megalocarpae suball. nova (Table 5 in Appendix)

    Diagnostic taxa: Craibia brownii, Markhamia lutea, Croton alienus, Rothmannia urcelliformis.

    Holotype: Crotono megalocarpae-Brachylaenetum huillensis Bussmann et Beck 1995 (Table 3 in

    Appendix)

    The Karura forest and one Oloolua forest (1613-1945 m) were characterized by Craibia

    brownii, Markhamia lutea, Croton alienus and Rothmannia urcelliformis, and lacking Schrebera

    alata, Turraea mombassana and Gnidia subcordata (diagnostic species of Schreberenion alatae).

    These diagnostic species are characteristic at Ngong, Oloolua and the Loita Hills. Karura is a quite

    undulating, hilly area, with more rivers than at Ngong. Two associations are distinguished: Crotono

    megalocarpae-Brachylaenetum huillensis Bussmann et Beck 1995 and Markhamio luteae-

    Veprisetum trichocarpae ass. nova.

    1-1-2-1-1. Markhamio luteae-Veprisetum trichocarpae ass. nova (Table 5 in Appendix)

    Diagnostic taxa: Craibia brownii, Markhamia lutea, Croton alienus, Rothmannia urcelliformis.

    Holotype: running no. 7 (Ken 81, 1710 m)

    This association is typical for Karura Forest. In most forests in Karura, Vepris trichocarpa,

    Croton megalocarpus, Rawsonia lucida and Strychnos usambarensis are dominant in the canopy

    layer. The forest height is 25-40 m. Areas of natural forest are well kept in the Karura Forests.

    Markhamio luteae-Veprisetum trichocarpae is classified into three subassociations: allophyletosum

    rubifoliae, warburgietosum ugandensis,and newtonietosum buchananii.

    1-1-2-1-1-1. allophyletosum rubifoliae subass. nova (Table 5 in Appendix)

    Diagnostic taxa: Allophylus rubifolius, Hippocratea africana, Crotalaria goodiiformis.

    Holotype: running no. 1 (201308 Kar-1, 1676 m)

    The height of this forest is 30-40 m, and the canopy is open (cover 50-65%). The

    undercanopy layer is of typical height at 20-22 m, and its cover is 70-80%. Only the forest of

    running no. 5 has higher values i.e., 23 m height and 80% canopy cover. This subassociation occurs

    on relatively flat areas. Cover of the forest floor is 20%, which is higher than in other

    subassociations.

    1-1-2-1-1-2. warburgietosum ugandensis subass. nova (Table 5 in Appendix)

    Diagnostic taxa: Warburgia ugandensis, Vepris hanangensis var. unifoliata, Ochna ovata,

    Allophylus ferruginus, Cissus oliveri, Vitex strickeri

    Holotype: running no. 8 (Ken-9, 1669 m)

    This subassociation is also tall forest, at 25-35 m, and has canopy cover of 60-80%. It occurs

    on relatively flat topography.

    1-1-2-1-1-3. newtonietosum buchananii subass. nova (Table 5 in Appendix)

    Diagnostic taxa: Newtonia buchananii, Uvariodendron anisatum

  • FOREST TYPES AND BIODIVERSITY AROUND THE GREAT RIFT VALLEY… 149

    Holotype: running no. 19 (Ken-8, 1887 m)

    This subassociation is characterized by the dominance of Newtonia buchananii in the

    canopy layer, which is 22-32 m high. Canopy cover is also high, at 75-80%. This forest type occurs

    from flat areas to steep slopes of 20-28% and at the highest elevations (1657-1945 m) of the three

    subassociations.

    1-1-2-2. Schreberenion alatae suball. nova (Table 3 in Appendix)

    Diagnostic taxa: Schrebera alata, Turraea mombassana, Gnidia subcordata

    Holotype: Euphorbio candelabrae-Oleetum africanae ass. nova

    One diagnostic species, the deciduous Schrebera alata (Oleaceae), occurs in open woodlands

    and drier forests. Forests at Ngong, Oloolua and the Loita Hills are classified into this suballiance.

    The field data were obtained at 1731-2024 m. This suballiance occurs in the southernmost hills.

    Two associations and one community are distinguished in this suballiance: Elaeodendro

    buchananii-Brachylaenetum huillensis Bussman et Beck 1995, Euphorbio candelabrae-Oleetum

    africanae ass. nova, and Canthium keniense-Brachylaena huillensis community.

    1-1-2-2-1. Euphorbio candelabrae-Oleetum africanae ass. nova (Table 6 in Appendix)

    Diagnostic taxa: Scolopia zeyheri, Scolopia theifolia, Euphorbia candelabrum, Abutilon longicuspe,

    Justicia anagalloides, Pappea capensis, Tinnea aethiopica, Tridactyle furcistipes, Maerua triphylla.

    Holotype: running no. 3 (Ken-225, 2189 m)

    Euphorbio candelabrae-Oleetum africanae has a unique physiognomy, with tall Euphorbia

    candelabrum. The height of forests is mostly 12-20 m, sometimes 27 m. Canopy cover is 25-60%,

    i.e. relatively open. This forest association occurs in the Loita Hills and on slopes of 5-20%.

    1-1-3. Mayteno undatae-Veprision simplicifoliae all. nova prov. (Table 3 in Appendix)

    Diagnostic taxa: Ehretia cymosa, Maytenus undata, Vangueria infausta, Fagaropsis angolensis

    Holotype: Crotono megalocarpae-Tecletum simplicifoliae Bussmann 2002

    This alliance is composed of two community types: Crotono megalocarpae-Tecletum

    simplicifoliae Bussmann 2002 and Desmodium repandum-Fagaropsis angolensis community. This

    alliance occurs widely in Mukogodo Forest, the Ngong Hills Plain, Karissia Hills, Bawa, Aberdare

    National Park, the Mattews Range and Loita Naimena Eukiyo Forest. This alliance includes

    Podocarpus falcata forests, Juniperus forests, and also evergreen broad-leaved forests.

    II. Afromontane forests

    White (1983) described Afromontane forests as occurring between 1200 and 2500 m, from

    southern Ethiopia to Malawai. Our data in Kenya are from 2100 m to 2600 m, where evergreen

    moist forests are described as the class Ocotetea.

    2. Ocotetea usambaraensis Bussmann et Beck 1995, Ocotetalia usambaraensis Bussmann et Beck

    1995

    Diagnostic taxa: Tabernaemontana stapfiana, Syzygium guineense, Ocotea usambarensis, Peddiea

    fischeri, Strombosia scheffleri, Panicum calvum, Piper capense, Schefflera volkensii, Xymalos

    monospora, Adenia gummifera.

    Holotype: Macarangion kilimandscharicae Bussmann et Beck 1995

  • 150 K. FUJIWARA, T. FURUKAWA, S. KIBOI, S. MATHENGE, P. MUTISO,

    H. HAYASHI, SHIN-ICHI MEGURO

    This class was described as including two orders by Bussmann and Beck [5], but diagnostic

    species of the two orders were not clear.

    2-1-1. Dracaena afromontana-Cassipourea malosana community (Table 3 in Appendix)

    Differential taxa: Dracaena afromontana, Acanthus eminens, Pavetta elliottii, Mimulopsis solmsii,

    Dicliptera laxata, Vepris (Teclea) nobilis, Simirestis goetzei, Canthium oligocarpum, Stombosia

    scheffleri, Bersama abyssinica.

    Holotype: running no. 16

    The new community type was recorded from the so-called Mau area that included the

    Tinderet Forest (Nandi Hills), Nyangores Block (Narok), Nyangores Forest Station (Narok) and

    Chemususu Lembus Forest (Eldama Ravine). Forests from the western hills of the Great Rift Valley

    are not surveyed well. Kakamega Forest is always studied and reported. The data were gathered

    from Nkareta Forest, Tinderet Forest, the Nyangores Block, Transmara Forest, Nyangores Forest

    and Chemususu Lembus Forest. This community occurs in the western hill area, at 2125-2318 m.

    The forest is tall (28-55 m) and canopy cover is 60-80%. It occurs on gentle slopes or almost flat

    topography.

    2-1-2. Macarangion kilimandscharicae Bussmann et Beck 1995

    Diagnostic taxa: Begonia meyeri-johannis, Lasianthus kilimandscharicus, Neoboutonia macrocalyx,

    Macaranga kilimandscharica, Plectranthus luteus.

    Holotype: Macarangetum kilimandscharicae by Bussmann et Beck 1995

    Macarangion kilimandscharicae Bussmann et Beck 1995 was described through certain

    diagnostic taxa. In particular, the Macarangetum kilimandscharicae sensu lato by Bussmann et Beck

    1995 was described from Mt. Kenya. Those authors, though, made too many associations based on

    herb species in Kenya, and these should be reorganized. Therefore, it seems that the unit by

    Bussmann [4] needs to be reconsidered (Table 3 in Appendix). Another forest type was recorded

    from Kieni Forest, Ragati Forest and Kiandongoro Forest and identified as Psychotria

    fractinervata-Macaranga kilimandscharica community, with the following differential species:

    Carex monostachya, Pauridiantha paucinervis, Keetia gueinzii, Psychotria fractinervata, Impatiens

    fischeri, Urera hypselodendron and Psychotria riparia. Podocarpo latifolii-Sinarundinarietum

    alpinae Bussmann et Beck 1995 does belong to Sinarundinarietea alpinae Bussmann et Beck 1995.

    It does not have character species of Ocotetea usambarensis or Juniperetea procerae, but common

    species do occur. It needs more consideration.

    III. Dry montane forests

    These forest were assigned to Juniperetea procerae Bussmann et Beck 1995 and

    Juniperetalia procerae Bussmann et Beck 1995.

    Diagnostic taxa: Juniperus procera, Stipa dregeana, Isoglossa gregorii, Ehrharta erecta.

    Holotype: Juniperion procerae Bussmann et Beck 1995

    3-1-1 Juniperion procerae Bussmann et Beck 1995 (Table 3 in Appendix)

    Diagnostic taxa: Geranium arabicum, Rhamnus prinoides, Rubus volkensii, Myrsine africana,

    Brachypodium flexum.

    Holotype: Myrsino africanae-Juniperetum procerae Bussmann et Beck 1995

  • FOREST TYPES AND BIODIVERSITY AROUND THE GREAT RIFT VALLEY… 151

    Six associations on Mt. Kenya were described in this alliance, but only three associations

    have clear diagnoses: Podocarpo latifolii-Cassipouretum malosanae Bussmann et Beck 1995,

    Myrsino africanae-Juniperetum procerae Bussmann 1995, and Faulio salignae-Ilicietum mitis

    Bussmann 2002 (Table 3 in Appendix). These occur on Mt. Kenya and Mt. Nyiro, and at Ngare,

    Porror, and Lorogi.

    3-1-2 Cassipourion malosanae Bussmann et Beck 1995

    Diagnostic taxa: Cassipourea malosana, Olea capensis ssp. hochstetteri, Lepidotrichilia volkensii,

    Pilea usambarensis

    Holotype: Tecleo nobilis-Tecletum simplicifoliae Bussmann et Beck 1995

    This plant association is summarized in this alliance from the Loita Hills, Ndare Ngare,

    Porror, and Lorogi.

    Discussion and Conclusions

    There are not enough climate data in Kenya, and Bussmann and Beck [5] and Bussmann [4]

    did not include detailed environmental data. The data used herein, however, do cover the whole

    area. Therefore we can discuss forest types and environmental factors based on DCCA results (see

    Figure 2).

    1. Characteristics of forest-type distribution along the Great Rift Valley

    The forests of Kenya and other parts of eastern Africa were described by [42, 35, 5, 4, 25].

    White [42] described the forests of Africa especially well. Tropical dry forest at Karura and the

    Ngong forest in Nairobi were reported by [24, 16, 11, 12], as well as by [4]. Bytebier and Bussmann

    [7] reported vegetation types and provided a taxa checklist for Mt. Nyiru. Githae [14] provided a

    botanical inventory and reported on diversity at Mt. Marsabit, at least partially. Hemp [17] reported

    on the vegetation of Mt. Kilimanjaro. Vegetation zonation is similar to that of Mt. Kenya except

    that there is no bamboo on Kilimanjaro. People use the forest in several ways [18, 19, 26, 33].

    Therefore, natural forests are now only remnants. Regeneration and succession in tropical dry forest

    and Afromontane forest have also been studied [3, 21, 34, 38].

    According to the DCCA results, the longitude, latitude, elevation, slope, aspect,

    precipitation, number of wetter and drier seasons, and coldest month temperature are major

    environmental factors. The forests of Marsabit and the Mattews Range are located the farthest east

    and north. These (running nos. 2 & 3 in Table 1, in Appendix) are ‘island forests’ in the desert [cf.

    4, 6, 13, 15, 20, 28, 29, 41, 42]. These are mostly tropical dry forests and are different from forests

    of the Nairobi area (nos. 5-10: Brachylaenion huillensis). Tropical dry forests are characterized by

    lower annual precipitation (see Figure 1), low wet-season precipitation, gentle slopes, southern

    aspects and southern latitude in the Nairobi area (nos. 4-11). Another type of tropical dry forest,

    without Brachylaenion huillensis species, is characterized by more western locations (hilly

    topography), more dry-season precipitation, low coldest mean temperature, just one (but a longer)

    dry season each year, and northern latitude, except for the Loita Hills (nos. 12-15). Afromontane

    forests (Ocotetea usambarensis and Juniperetea procerae) are characterized by the lowest

    temperatures and high elevation (nos. 16-20). The highest-elevation forest in our relevé set was on

    Mt. Aberdare, at 3030 m (no. 21). It is a unique Podocarpus forest with Arundinaria, no dry-forest

    species, and some moist-forest tree species. The Afromontane forests occur in relatively inland

    mountains. These sites are at high elevation, where the coldest-month temperature is close to 0ºC

    (nos. 16, 20, 21).

  • 152 K. FUJIWARA, T. FURUKAWA, S. KIBOI, S. MATHENGE, P. MUTISO,

    H. HAYASHI, SHIN-ICHI MEGURO

    Fig. 2: DCCA (De-trended Canonical Correspondence Analysis) of forest types along the Great Rift Valley

    Top: The numbers 2-21 are the running numbers of Table 3 (see Appendix). Only those vegetation samples with

    environmental data (see Table 2 in Appendix)were analyzed by the DCCA.

  • FOREST TYPES AND BIODIVERSITY AROUND THE GREAT RIFT VALLEY… 153

    Caption of Figure 2 (continued)

    Oval A represents the Tropical Dry Forests in the main text: Schoenoxiphio lehmannii-Crotonietalia megalocarpae. Its

    subsets correspond to subsections in the main text: A-1: Strychnoso mitis-Diospyrosion abyssinicae; A-2:

    Brachylaenion huillensis; A-3: Mayteno undatae-Veprision simplicifoliae. Ovals B and C represent Afromontane

    Forests (see main text): Ocotetea usambarensis (B) and Podocarpo latifolii-Sinarundinarietum alpinae (C).

    Bottom: Results of the DCCA, based on the following environmental factors: ELE = elevation, RSP = precipitation sum

    of the rainy season, AP = annual precipitation, ASP = aspect, AT = annual mean temperature, CMT = mean temperature

    of coldest month, LON = Longitude, DPS = precipitation sum of the dry season, FDS = number of dry periods per year,

    LAT = latitude, SLO = slope, and CT1 = Cover percentage of the canopy layer (T1).

    2. Data differences from Bussmann and Beck 1995 and Bussmann 2002.

    Detailed phytosociological data for forests in Kenya were reported by [5, 4]. Their data were

    integrated with the present authors' data in Table 1 (see Appendix). It is curious that data from

    Bussmann suggest independent association types, such as at Karura and Ngong, Mt. Marsabit, Mt.

    Kulal, the Loita Hills and the Mattews Range. The two classes of evergreen and semi-evergreen

    forest, plus the Sinoarundinarietea (bamboo class) and Hagenietea on Mt. Kenya, were classified

    fundamentally. Four classes, five orders, 10 alliances and 41 associations were reported from Mt.

    Kenya [5]. These data were integrated with the new data, which allowed the distinction of one new

    order and one new class (Schoenoxiphio lehmannii-Crotonietalia megalocarpi and Schoenoxiphio

    lehmannii-Crotonietea megalocarpi) circumscribed to tropical dry forests. These did belong to the

    Juniperetea procerae before, but common species were not clear. The Ocotetea usambarensis

    Bussmann et Beck 1995 was difficult to order. Here the authors made the new units but leave the

    question for later discussion, because many associations are classified by herbs. Two orders and

    four alliances could not be clarified. Here, new data from Mau Mt. suggested a new association and

    alliance.

    In this paper three classes were discussed. Unfortunately, new relevés from Mt. Kenya were

    not obtained, partly since most forests had been destroyed. We guess that the forests would have

    been in better condition in 1995, when Bussmann and Beck made their vegetation samples.

    Macarangion kilimandscharicae constitutes the major part of the Afromontane forests on Mt.

    Kenya. Three classes were followed by Ocotetea usambarensis and Juniperetea procerae that

    Bussmann and Beck [5] described in 1995. There are several tree species common in Kenyan

    evergreen and semi-evergreen forests. The situation of these species needs to be discussed. The

    authors will do more vegetation samples and record other natural Kenyan forests in the future.

    Acknowledgements: We wish to express our thanks to foresters of the Forest Department of the Kenyan Forest

    Service; also to Mr. S. Kage, IFCM, for the whole study period, and to former staff members of the Green Belt

    Movement (2006-2007); to Graduate School students of the former Faculty of Vegetation Science of Yokohama

    National University (2006-2009) and to Dr. L. Borghesio (2007, 2008); and to Prof. E.O. Box (2008) (University of

    Georgia), Mr. Chebee (2010-2013), Mr. C. Mutiso (2014) (University of Nairobi), and Mrs. Atsuko Harada (in 2012)

    for helping our field survey. We also thank Dr. You Hai-Mei (Jiangsu Normal University, in China) for analyzing the

    data by DCCA; and Mrs. Masami Sugita and Dr. Chie Itow for organizing the data and making the figures and tables

    for this manuscript. We are grateful to Prof. E.O. Box for English editing of this manuscript and for providing the

    climate data for Kenya. This survey was funded by a grant from HIOKI Co. Ltd. for the project of restoration of natural

    forests in Kenya.

  • 154 K. FUJIWARA, T. FURUKAWA, S. KIBOI, S. MATHENGE, P. MUTISO,

    H. HAYASHI, SHIN-ICHI MEGURO

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    TIPURILE DE PĂDURE ȘI BIODIVERSITATEA DIN ZONA VĂII MARELUI RIFT DIN KENYA

    (Rezumat)

    Pădurile din Valea Marelui Rift din Kenya sunt unice, ca insule verzi în deșert și se extind din sudul Etiopiei

    până în Tanzania. Aceste păduri le includ pe cele Afromontane și tropical-uscate, studiate în Kenya de către Bussmann

    și Beck, iar în sudul Etiopiei de Bussmann. Au fost descrise două clase taxonomice (Ocotetea usambarensis în climatul

  • 156 K. FUJIWARA, T. FURUKAWA, S. KIBOI, S. MATHENGE, P. MUTISO,

    H. HAYASHI, SHIN-ICHI MEGURO umed și Juniperetea procerae în locurile uscate) anterior studiilor noastre care s-au desfășurat pe părțile estică și vestică

    ale Văii Marelui Rift, în perioada 2006-2014, iar rezultatele au fost comparate cu cele ale lui Bussmann din 1990.

    Această lucrare prezintă următoarele rezultate noi: 1) Pădurile rămân pe partea vestică și la peste 2000 m și sunt

    edificate de Ocotea kenyensis (Lauraceae); acestea sunt diferite de cele Afromontane umede din zonele mai estice

    (Ocotetea usambarensis) lângă Valea Riftului. Din martie până în aprilie și din august în septembrie sunt 4 luni cu

    precipitații de peste 100 mm. 2) La altitudini de 1300-2200 m, pe partea estică, sunt pădurile uscate unice, edificate de

    Brachylaena huilliensis (Compositae), Warburgia ugandensis (Canellaceae), Elaeodendron buchananii (Celastraceae),

    Vepris trichocarpa (Rutaceae), Calodendrum capense etc. Unele dintre aceste specii sunt unice și se întind până în

    pădurile tropicale din Africa de Sud. 3) Pădurile de altitudine ridicată din zonele estice sunt edificate de Podocarpus

    latifolius și Juniperus procera, care aparțin la Juniperetea procerae. Pădurile de Podocarpus latifolius apar în zonele

    vestice, dar sunt diferite de cele din zonele estice. 4) Pădurile din munții nordici izolați Marsabit și Kulal sunt diferite;

    nu există Podocarpus sau Juniperus, iar umiditatea ridicată permite apariția specie Usnea usneoides la Marsabit. 5)

    Zona împădurită a Kenyei este teoretic de 12%, iar pădurile naturale rămase ocupă doar 1,9%. Această suprafață mică

    permite eșalonarea unor tipuri diferite de păduri de-a lungul gradientului de umiditate. 6) Numeroase specii lemnoase și

    de ferigi din pădurile Afromontane sunt comune și în pădurile tropicale montane și subtropicale din Asia, inclusiv specii

    de Podocarpus, Juniperus, Myrsine, Rapanea, Psychotria, Rumohra, Asplenium, Pteris și Dryopteris. 7) Clasificările

    făcute de noi și cea a lui Bussmann nu sunt identice la nivel de asociație sau comunitate. De aici și întrebarea dacă

    informațiile din 1990 au fost colectate din zone diferite, sau dacă, o parte din vegetație a dispărut datorită impactului

    uman sau al faunei sălbatice.

    Received: 17.11.2014; Accepted: 27.11.2014.

  • FOREST TYPES AND BIODIVERSITY AROUND THE GREAT RIFT VALLEY… 157

    Tab

    le 1

    : Site

    loca

    tions

    and

    cor

    resp

    onde

    nce

    with

    Fig

    ure

    2 an

    d T

    able

    3.

    Ref

    . no.

    in F

    ig. 2

    Run

    ning

    no.

    in T

    able

    3L

    ocat

    ion

    -1

    Mt K

    ulal

    22

    Mar

    sabi

    t Nat

    iona

    l Par

    k3

    -M

    t. N

    yiro

    43,

    14

    Mat

    thew

    s Ran

    ge (K

    itich

    i Cam

    p)5

    17, 1

    8, 2

    6M

    atth

    ews R

    ange

    (Rap

    a-O

    loik

    Mat

    thew

    s Ran

    ge, O

    rkai

    la, L

    onda

    dapo

    )6

    18K

    aris

    sia

    Hill

    s (Sa

    anat

    a)7

    15N

    abol

    o Ro

    ck, B

    awa,

    Kar

    issi

    a H

    ills

    815

    , 16,

    18

    Kar

    issi

    a H

    ills (

    Nab

    olo

    Rock

    , Lol

    geriy

    io, N

    gabo

    lo)

    912

    Kar

    issi

    a H

    ills (

    Lem

    etek

    i)10

    22, 2

    7, 2

    9N

    gaia

    Hill

    s, N

    dare

    Nga

    re, P

    orro

    r, Lo

    rogh

    i, N

    yam

    beni

    Hill

    s11

    13M

    ukog

    odo

    Fore

    st, D

    ol D

    ol v

    illag

    e, K

    opia

    Val

    ley

    (Kur

    i Kur

    i Vill

    age)

    , Muk

    ogod

    o Fo

    rest

    , Dol

    Dol

    vill

    age

    1220

    Che

    mus

    usu

    Lem

    bus F

    ores

    t(El

    dam

    a Ra

    vine)

    1320

    Tind

    eret

    For

    est (

    Nan

    di H

    ills)

    1428

    Abe

    rdar

    e N

    atio

    nal P

    ark

    1521

    , 23,

    24,

    30,

    31

    Raga

    ti Fo

    rest

    1624

    Kia

    ndon

    goro

    For

    est

    1720

    Tran

    smar

    a Fo

    rest

    (Nya

    ngor

    es F

    ores

    t, C

    ham

    anen

    airo

    tia F

    ores

    t Sta

    tion)

    1824

    Kie

    ni F

    ores

    t, K

    iam

    bu D

    istri

    ct19

    19N

    kare

    ta F

    ores

    t20

    22G

    akoe

    215,

    6, 8

    Nai

    robi

    arb

    oret

    um, K

    arur

    a Fo

    rest

    , Kar

    ura

    fore

    st (N

    airo

    bi)

    -7,

    9, 1

    0, 2

    5N

    gong

    Roa

    d Fo

    rest

    San

    ctua

    ry (N

    airo

    bi);

    Ngo

    ng R

    oad

    Fore

    st, N

    airo

    bi (r

    eplic

    ate

    of K

    en-1

    2), N

    gong

    Roa

    d Fo

    rest

    , Nai

    robi

    ; Ngo

    ng F

    ores

    t, N

    airo

    bi; O

    lool

    ua F

    ores

    t (N

    gong

    Hill

    s For

    est S

    tatio

    n), N

    airo

    bi; O

    lool

    ua

    (Mus

    eum

    For

    est),

    Nai

    robi

    ; Olo

    olua

    For

    est,

    Nai

    robi

    -13

    ,25

    Ngo

    ng p

    lain

    s (w

    est o

    f Ngo

    ng H

    ills i

    nsid

    e th

    e G

    reat

    Rift

    Val

    ley)

    Ngo

    ng H

    ills P

    lain

    (Rift

    Val

    ley)

    Ngo

    ng

    Hill

    s, N

    airo

    bi

    -4,

    11,

    18

    Loita

    For

    est,

    Olo

    osuy

    ian

    (Loi

    ta N

    aim

    ena

    Euki

    yo F

    ores

    t) Lo

    ita F

    ores

    t, N

    esuk

    ari L

    oita

    For

    est,

    Nko

    pon

    Loita

    Fo

    rest

    (Loi

    ta N

    aim

    ena

    Euki

    yo F

    ores

    t)

    APP

    EN

    DIX

  • 158 K. FUJIWARA, T. FURUKAWA, S. KIBOI, S. MATHENGE, P. MUTISO, H. HAYASHI, SHIN-ICHI MEGURO

    Table no.Running no.

    Relevé no.

    Dat

    eL

    ocat

    ion

    T1 height (m)

    T2 height (m)

    S1 height (m)S2 height (m)

    H height (m)

    T1 cover (%)

    T2 cover (%)S1 cover (%)

    S2 cover (%)

    H cover (%)

    Elevation (m)

    Aspect

    Slope (°)

    Area (m²)

    Lat

    itude

    Lon

    gitu

    de

    1K

    l 106

    /02/

    2014

    Mt K

    ulal

    (Mar

    sabi

    t)42

    2216

    70.

    570

    3030

    2050

    1916

    S80E

    1530

    ×35

    N02

    °40'

    00.1

    "E3

    6°56

    '29.

    0"2

    Kl 2

    06/0

    2/20

    14M

    t Kul

    al (M

    arsa

    bit)

    3622

    145

    0.3

    6050

    2520

    6019

    18S2

    0E25

    25×3

    0N

    02°3

    9'58

    .8"

    E36°

    56'3

    0.0"

    3K

    l 306

    /02/

    2014

    Mt K

    ulal

    (Mar

    sabi

    t)38

    2416

    60.

    380

    4030

    2525

    1946

    S15

    25×3

    0N

    02°3

    9'59

    .2"

    E36°

    56'3

    2.5"

    4K

    l 407

    /02/

    2014

    Mt K

    ulal

    (Mar

    sabi

    t)27

    1912

    70.

    460

    3030

    5020

    1835

    S50W

    530

    ×40

    N02

    °40'

    06.6

    "E3

    6°56

    '45.

    4"5

    Kl 5

    07/0

    2/20

    14M

    t Kul

    al (M

    arsa

    bit)

    3322

    146

    0.5

    5540

    3060

    5019

    75S6

    0E20

    30×3

    5N

    02°4

    0'07

    .9"

    E36°

    56'3

    6.3"

    6K

    l 607

    /02/

    2014

    Mt K

    ulal

    (Mar

    sabi

    t)30

    2215

    70.

    440

    6030

    2550

    1940

    S10E

    1030

    ×30

    N02

    °40'

    03.2

    "E3

    6°56

    '26.

    5"7

    Mar

    _101

    /02/

    2014

    Mar

    sabi

    t Nat

    iona

    l Par

    k 35

    2010

    50.

    760

    5075

    401

    1337

    S75E

    1330

    ×35

    N02

    °15'

    45.0

    "E3

    7°55

    '50.

    6"8

    Mar

    _202

    /02/

    2014

    Mar

    sabi

    t Nat

    iona

    l Par

    k35

    2414

    60.

    475

    2010

    4530

    1540

    S40E

    1530

    ×30

    N02

    °18'

    53.5

    "E3

    7°58

    '22.

    1"9

    Mar

    _302

    /02/

    2014

    Mar

    sabi

    t Nat

    iona

    l Par

    k38

    2214

    60.

    570

    3025

    3025

    1539

    S35E

    1830

    ×25

    N02

    °18'

    19.8

    "E3

    7°57

    '42.

    7"10

    Mar

    _402

    /02/

    2014

    Mar

    sabi

    t Nat

    iona

    l Par

    k27

    2312

    60.

    360

    3050

    1025

    1520

    S35E

    1030

    ×25

    N02

    °18'

    08.1

    "E3

    7°57

    '32.

    8"11

    Mar

    _503

    /02/

    2014

    Mar

    sabi

    t Nat

    iona

    l Par

    k37

    2012

    30.

    350

    5070

    153

    1410

    L0

    35×3

    5N

    02°1

    6'16

    .0"

    E37°

    55'4

    7.5"

    12M

    ar_6

    03/0

    2/20

    14M

    arsa

    bit N

    atio

    nal P

    ark

    3422

    144

    0.5

    6040

    7025

    313

    92S7

    5E5

    25×4

    0N

    02°1

    5'12

    .3"

    E37°

    56'0

    2.4"

    13M

    ar_7

    03/0

    2/20

    14M

    arsa

    bit N

    atio

    nal P

    ark

    3724

    187

    0.6

    6060

    5020

    512

    30S5

    5W3

    25×4

    0N

    02°1

    4'39

    .3"

    E37°

    55'5

    5.9"

    14M

    ar_8

    04/0

    2/20

    14M

    arsa

    bit N

    atio

    nal P

    ark

    3723

    178

    0.3

    7050

    6020

    513

    00L

    035

    ×35

    N02

    °18'

    35.3

    "E3

    7°59

    '41.

    9"15

    Mar

    _904

    /02/

    2014

    Mar

    sabi

    t Nat

    iona

    l Par

    k37

    2314

    70.

    480

    6030

    2010

    1410

    S10W

    535

    ×50

    N02

    °18'

    34.5

    "E3

    7°58

    '47.

    3"16

    Mar

    _10

    04/0

    2/20

    14M

    arsa

    bit N

    atio

    nal P

    ark

    3823

    164

    0.5

    5050

    4020

    5012

    96S5

    0E3

    10×2

    0N

    02°1

    8'34

    .7"

    E38°

    00'0

    1.1"

    17M

    ar_1

    104

    /02/

    2014

    Mar

    sabi

    t Nat

    iona

    l Par

    k32

    2214

    60.

    260

    5040

    205

    1390

    N50

    E20

    30×4

    0N

    02°1

    6'57

    .9"

    E37°

    57'1

    5.1"

    18K

    en-8

    316

    /09/

    2007

    Mat

    tew

    s Ran

    ge, K

    itich

    i Cam

    p21

    .414

    -4

    0.7

    7060

    -25

    3014

    35S1

    0W10

    20x2

    0N

    01˚1

    5'10

    .8"

    E37˚

    17'3

    0.7"

    19K

    en-8

    416

    /09/

    2007

    Mat

    tew

    s Ran

    ge, K

    itich

    i Cam

    p29

    .116

    -4

    0.4

    7575

    -30

    2514

    45N

    20W

    1625

    x25

    N01˚1

    5'13

    .6"

    E37˚

    17'2

    8.0"

    20K

    en-8

    516

    /09/

    2007

    Mat

    tew

    s Ran

    ge, K

    itich

    i Cam

    p33

    .516

    6-

    0.5

    8040

    55-

    1014

    40N

    80W

    1020

    x25

    N01˚1

    5'11

    .7"

    E37˚

    17'3

    7.5"

    21K

    en-8

    616

    /09/

    2007

    Mat

    tew

    s Ran

    ge, K

    itich

    i Cam

    p33

    .618

    6-

    0.5

    7550

    35-

    514

    75N

    85W

    2112

    x20

    N01˚1

    5'08

    .7"

    E37˚

    17'4

    3.2"

    22K

    en-8

    717

    /09/

    2007

    Mat

    tew

    s Ran

    ge, K

    itich

    i Cam

    p27

    .116

    6-

    0.6

    7560

    35-

    2014

    35S8

    0E12

    20x2

    0N

    01˚1

    4'45

    .7"

    E37˚

    17'3

    0.0"

    23K

    en-8

    817

    /09/

    2007

    Mat

    tew

    s Ran

    ge, K

    itich

    i Cam

    p27

    127

    -1.

    565

    5025

    -80

    1445

    N40

    E12

    25x3

    0N

    01˚1

    4'47

    .4"

    E37˚

    17'2

    6.8"

    24K

    en-8

    917

    /09/

    2007

    Mat

    tew

    s Ran

    ge, K

    itich

    i Cam

    p30

    .620

    6-

    0.6

    7055

    60-

    1514

    20N

    60E

    520

    x25

    N01˚1

    4'47

    .5"

    E37˚

    17'2

    7.0"

    25K

    en-9

    017

    /09/

    2007

    Mat

    tew

    s Ran

    ge, K

    itich

    i Cam

    p32

    176

    -0.

    570

    5035

    -25

    1425

    S80W

    623

    x18

    N01˚1

    4'48

    .1"

    E37˚

    17'3

    6.2"

    Table 4Tab

    le 2

    : Rel

    evé

    data

    in T

    able

    s 4-6

    (L: l

    evel

    asp

    ect;

    Asp

    ect b

    y cl

    inom

    eter

    : Kam

    iyam

    a Se

    isak

    usho

    211

    -000

    8; L

    atitu

    de &

    long

    itude

    by

    GPS

    : GA

    RM

    IN

    eTre

    x30J

    ).

  • FOREST TYPES AND BIODIVERSITY AROUND THE GREAT RIFT VALLEY… 159

    26K

    en-9

    117

    /09/

    2007

    Mat

    tew

    s Ran

    ge, K

    itich

    i Cam

    p36

    206

    -0.

    580

    7525

    -10

    1440

    N80

    W16

    30x2

    0N

    01˚1

    4'51

    .6"

    E37˚

    17'4

    1.1"

    27K

    en-1

    0622

    /09/

    2007

    Mat

    tew

    s Ran

    ge, K

    itich

    i Cam

    p28

    165

    -0.

    550

    8035

    -10

    2043

    S50W

    2018

    x20

    N01˚1

    4'53

    .9"

    E37˚

    17'3

    7.7"

    28K

    en-1

    0722

    /09/

    2007

    Mat

    tew

    s Ran

    ge, K

    itich

    i Cam

    p33

    186

    -0.

    660

    7030

    -5

    1425

    S50W

    1520

    x20

    N01˚1

    4'38

    .9"

    E37˚

    17'4

    5.5"

    29K

    en-1

    0822

    /09/

    2007

    Mat

    tew

    s Ran

    ge, K

    itich

    i Cam

    p30

    .522

    7-

    0.5

    5070

    40-

    1014

    40S8

    0W18

    25x2

    5N

    01˚1

    4'41

    .5"

    E37˚

    17'4

    6.7"

    30K

    en-1

    0922

    /09/

    2007

    Mat

    tew

    s Ran

    ge, K

    itich

    i Cam

    p25

    157

    20.

    650

    7035

    2025

    1475

    S50W

    2120

    x15

    N01˚1

    4'44

    .8"

    E37˚

    17'4

    8.4"

    1K

    ar 1

    17/0

    8/20

    13K

    arur

    a fo

    rest

    3522

    9-

    0.5

    6570

    25-

    2016

    76S6

    0W5

    15x2

    0S0

    1°14

    ’33.

    3"E3

    6°50

    '19.

    3"2

    Kar

    217

    /08/

    2013

    Kar

    ura

    fore

    st40

    206

    -0.

    460

    7025

    -20

    1674

    S50W

    530

    x30

    S01°

    14'3

    0.1"

    E36°

    50'1

    4.9"

    3K

    ar 3

    17/0

    8/20

    13K

    arur

    a fo

    rest

    3020

    8-

    0.5

    5075

    20-

    2017

    10L

    030

    x30

    S01°

    14'2

    1.1"

    E36°

    49'3

    9.3"

    4K

    ar 4

    17/0

    8/20

    13K

    arur

    a fo

    rest

    3520

    8-

    0.6

    6080

    25-

    2017

    13L

    020

    x30

    S01°

    14'1

    5.3"

    E36°

    49'3

    4.0"

    5K

    ar 5

    17/0

    8/20

    13K

    arur

    a fo

    rest

    2314

    -4

    0.8

    8030

    -35

    2017

    13L

    020

    x20

    S01°

    14'1

    2.1"

    E36°

    49'3

    2.4"

    6K

    en-2

    419

    /10/

    2006

    Kar

    ura

    Fore

    st26

    .315

    63

    0.4

    5060

    3020

    516

    92L

    021

    x23

    S01˚

    14'1

    9.0"

    E36˚

    49'2

    6.1"

    7K

    en-8

    102

    /05/

    2007

    Kar

    ura

    Fore

    st27

    .515

    5-

    0.6

    7070

    50-

    2017

    10N

    5E6

    15x3

    5S0

    1˚14

    '00"

    E36˚

    49'0

    0"8

    Ken

    -911

    /03/

    2006

    Kar

    ura

    Fore

    st32

    198

    30.

    640

    8510

    3010

    1613

    S40W

    530

    x35

    S01˚

    14'3

    1.3"

    E36˚

    50'1

    8.3"

    9K

    en-2

    319

    /10/

    2006

    Kar

    ura

    Fore

    st31

    156

    -0.

    580

    8090

    -3

    1669

    S20W

    321

    x21

    S01˚

    14'3

    6.2"

    E36˚

    50'2

    3.1"

    10K

    en-2

    519

    /10/

    2006

    Kar

    ura

    Fore

    st35

    .716

    6-

    0.3

    7050

    20-

    316

    70S2

    0W18

    18x2

    1S0

    1˚14

    '30.

    0"E3

    6˚49

    '31.

    1"11

    Ken

    -61

    24/0

    4/20

    07K

    arur

    a Fo

    rest

    27.2

    135

    -0.

    370

    6050

    -10

    1726

    L0

    35x1

    0S0

    1˚14

    '19.

    4"E3

    6˚49

    '40.

    3"12

    Ken

    -63

    24/0

    4/20

    07K

    arur

    a Fo

    rest

    27.2

    135

    -0.

    370

    6050

    -10

    1726

    L0

    35x1

    0S0

    1˚14

    '19.

    4"E3

    6˚49

    '40.

    3"13

    Ken

    -64

    26/0

    4/20

    07K

    arur

    a Fo

    rest

    2515

    3-

    0.6

    6075

    25-

    1016

    89N

    20W

    315

    x26

    S01˚

    14'3

    0.6"

    E36˚

    50'1

    3.2"

    14K

    en-7

    902

    /05/

    2007

    Kar

    ura

    Fore

    st25

    164

    -0.

    680

    7040

    -30

    1726

    L0

    25x2

    5S0

    1˚14

    '11.

    4"E3

    6˚49

    '39.

    0"15

    Ken

    -80

    02/0

    5/20

    07K

    arur

    a Fo

    rest

    2416

    5-

    0.5

    7060

    50-

    2517

    20N

    42E

    830

    x20

    S01˚

    14'2

    3.3"

    E36˚

    49'2

    5.8"

    16K

    en-2

    118

    /10/

    2006

    Kar

    ura

    Fore

    st31

    .514

    6-

    0.6

    7530

    30-

    6016

    45S2

    5E4

    20x2

    5S0

    1˚14

    '42.

    3"E3

    6˚50

    '27.

    7"17

    Ken

    -44

    19/0

    4/20

    07O

    lool

    ua F

    ores

    t26

    .315

    4-

    0.7

    8055

    40-

    3517

    80S1

    0E28

    36x1

    5S0

    1˚21

    '28.

    5"E3

    6˚42

    '41.

    1"18

    Ken

    -10

    11/0

    3/20

    06K

    arur

    a Fo

    rest

    2214

    74

    0.4

    7510

    510

    6016

    57N

    20E

    38x

    20S0

    1˚14

    '03.

    6"E3

    6˚49

    '32.

    9"19

    Ken

    -811

    /03/

    2006

    Kar

    ura

    Fore

    st35

    158

    30.

    685

    2520

    3510

    1887

    L0

    25x2

    5S0

    1˚14

    '35.

    7"E3

    6˚50

    '21.

    0"20

    Ken

    -22

    19/1

    0/20

    06K

    arur

    a Fo

    rest

    32.1

    164

    -0.

    380

    2580

    -35

    1945

    N25

    E20

    21x9

    S01˚

    14'4

    2.9"

    E36˚

    50'2

    6.0"

    1K

    en-2

    2026

    /09/

    2009

    Loita

    For

    est,

    Nko

    pon

    -12

    -6

    0.5

    -40

    -70

    519

    88S8

    0E5

    10x1

    5S0

    1˚50

    '07.

    0"E3

    5˚48

    '35.

    8"2

    Ken

    -221

    26/0

    9/20

    09Lo

    ita F

    ores

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    -225

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    Table 5 Table 6

  • 160 K. FUJIWARA, T. FURUKAWA, S. KIBOI, S. MATHENGE, P. MUTISO, H. HAYASHI, SHIN-ICHI MEGURO

    Run

    ning

    num

    ber

    12

    34

    56

    78

    910

    1112

    1314

    1516

    1718

    1920

    2122

    2324

    2526

    2728

    2930

    31E

    leva

    tion

    (m) /

    R

    efer

    ence

    (B &

    B

    1995

    = B

    ussm

    ann

    et

    Beck

    , 199

    5; B

    200

    2 =

    Buss

    man

    n, 2

    002)

    Layer

    1835-1940

    1230-1539

    1379-1561

    B et B 1995

    1674-1713

    1613~1726

    1657-1780

    B 2002

    1731-1959

    1828-2174

    1971-2189

    1803-1960

    1863-2154

    1930-2165

    2115-2277

    2109-2604

    B et B 1995

    B 2002

    B et B 1995

    2120-2460

    3080

    B 2002

    B et B 1995

    B et B 1995

    B 2002

    B 2002

    B 2002

    B 2002

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    Rel

    evé

    num

    ber

    611

    1310

    511

    430

    296

    65

    1512

    3221

    156

    4310

    37

    15

    126

    100

    4811

    2113

    146

    14N

    umbe

    r of

    spec

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    3627

    4249

    3738

    3852

    4343

    4736

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    48D

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    Tab

    le 3

    : Syn

    optic

    tabl

    e of

    fore

    sts c

    omm

    uniti

    es a

    long

    the

    Gre

    at R

    ift V

    alle

    y in

    Ken

    ya (T

    : Tre

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    : shr

    ub la

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    quen

    cy:

    V: 8

    1-10

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    1-80

    %; I

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    1-60

    %; I

    I: 2

    1-40

    %; I

    : 11-

    20%

    ; +: 6

    -10%

    ; r: <

    5%

    ).

  • FOREST TYPES AND BIODIVERSITY AROUND THE GREAT RIFT VALLEY… 161

    Pseu

    dera

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    HD

    orst

    enia

    bro

    wnii

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    ・・

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    gea

    abys

    sini

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    ・II

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    gnos

    tic sp

    ecie

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    roto

    no m

    egal

    ocar

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    chyl

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    is B

    ussm

    ann

    et B

    eck

    1995

    Vern

    onia

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  • 162 K. FUJIWARA, T. FURUKAWA, S. KIBOI, S. MATHENGE, P. MUTISO, H. HAYASHI, SHIN-ICHI MEGURO

    Vepr

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