DIVERSITY AND DISTRIBUTION OF THE AFROALPINE FLORA OF EASTERN
AFRICA WITH SPECIAL REFERENCE TO THE TAXONOMY OF THE GENUS
PENTASCHISTIS (POACEAE)
AHMED ABDIKADIR ABDI
(MSC)
REG. No. I84/10955/2007
A THESIS SUBMITTED IN FULFILLMENT OF THE REQUIREMENTS FOR THE
AWARD OF THE DEGREE OF DOCTOR OF PHILOSOPY IN THE SCHOOL OF
PURE AND APPLIED SCIENCES OF KENYATTA UNIVERSITY.
MAY, 2013
ii
DECLARATION
This thesis is my original work and has not been presented for a degree in any other
University or any other award.
Ahmed Abdikadir Abdi (MSc)
REG. No. 184/10955/2007
Signature:
Date
We confirm that the work reported in this thesis was carried out by the candidate under our
supervision.
1.
Prof. Leonard E. Newton, Kenyatta University, P.O. Box 43844−00100, Nairobi
Signature
2.
Date
Dr. Geoffrey Mwachala, East African Herbarium, National Museums of Kenya, P.O.
Box 40658−00100, Nairobi
Signature
Date
iii
DEDICATION
To AL–MIGHTY ALLAH whose Glory and Power sustains the great diversity of life and
the Universe!
iv
ACKNOWLEDGEMENT
I am grateful to the Norwegian Programme for development, Research & Higher Education
(NUFU) for funding the field work expenses and part of the tuition fees through AFROALP
II Project. I also sincerely thank the administrators and laboratory technologists of National
Agricultural Research Laboratories of Kenya Agricultural Research Institute particularly Dr
Gichuki, Dr Miano, Mr. Irungu, Mr. Mbogoh and Mr. Juma for their logistical and technical
support during laboratory work in their NARL−KARI laboratory. Thanks to all the wardens
and administrators of Bale and Simen Mountain National Park, Uganda Wildlife Authority,
Arusha and Meru National Park and Kenya Wildlife Service for permission to do field work
in their respective mountains. I sincerely appreciate assistance given to me by curators of
East African herbarium at Nairobi, National herbarium Addis Ababa University, Makerere
University herbarium, Uganda, and National herbarium in Arusha, Tanzania. I am very
grateful to all those who in one way or another contributed to the successful completion of
my field work, laboratory analysis and the overall work of this study.
v
ABSTRACT
The African tropic−alpine system, consists of a number of geographically separated
‘sky−islands’ and constitutes an excellent system in which to investigate the interaction
between the geographical and ecological components of differentiation. The mountain
systems harbouring the Afroalpine zone act as giant water towers that are source of almost all
rivers that are an important source of water for domestic, industrial and agricultural use. Due
to inaccessibility caused by extreme isolation and harsh conditions little research has been
done on the diversity, species richness and evenness among other aspects in the Afroalpine
zone. For effective conservation of these important zones it is important to understand their
ecology. and the conservation status of the species of these zones. The aim of this study was
to carry out ecological analysis of Afroalpine species in general and addresses species
delimitation in the genus Pentaschistis. The vegetation of Afroalpine zone of Mount Kenya,
Elgon, Kilimanjaro, Ruwenzori, Bale and Simen were sampled for ecological analysis and
taxonomic revision of the genus Pentaschistis. A total of 75 plots (100 × 100 m) from five
vegetation types were analysed using both univariate and multivariate analyses. From the
study a total of 46 families, 124 genera and 278 species were recorded from rock outcrops,
bogs, grassland, Dendrosenecio forest/woodland and Alchemilla vegetation communities. Six
families, namely Asteraceae, Poaceae, Caryophyllaceae, Lamiaceae, Cyperaceae and
Rosaceae constitute 56% of all the species sampled. Five communities were described from
the vegetation types studied: Carex−Deschampsia bogs, Festuca−Pentaschistis grassland
communities, Dendrosenecio−Alchemilla forest/woodland, Alchemilla communities and
Helichrysum−Festuca−Koeleria−Pentaschistis rock outcrop communities. The species
richness and diversity is higher in older mountains such as Mt. Simen, Bale and Elgon (over
20 MYBP) compared to younger mountains such as Mt. Kenya (3.5 MYBP), Kilimanjaro and
Rwenzori (less than 3 MYBP) mountains. Similarly Ethiopian Mountains were more species
rich and have higher diversity than mountains of East Africa. Simen was the most species
rich as well as exhibiting highest species diversity while Rwenzori was the most species poor
and with the least species diversity. The East Africa Mountains were similar both in their
species composition and community structure, as were the Ethiopian Mountains. Factors such
as the geographical distance, age, geological history, and position of the mountains along the
Rift Valley, climatic and edaphic factors are thought to be responsible for the observed
patterns. The five vegetation communities were recognizable from each other though rock
outcrop and grassland communities were barely separable (R< 0.25). The rest of the
vegetation communities were overlapping but separable especially between bogs vs.
Dendrosenecio (R > 0.5). The mainly southern African grass genus Pentaschistis was
represented on all Afro−alpine mountain systems but due to the complex ecological and
geographical variation patterns, the number of species recognized is widely disputed. A
classification based on a well–supported evolutionary hypothesis for the genus is necessary.
In order to obtain this, morphological studies of both natural populations and herbarium
specimens have been conducted during this study. A total of 38 characters were used to carry
out phenetic analysis of 37 specimens from nine taxa. The results from morphological
observation as well as the cluster and principal component analysis produced two main
groupings viz. (1) the two widely distributed species P. borussica and the P. pictigluma
species complex, and (2) two narrow endemics P. dolichochaeta and P. chrysurus. Similarly,
phylogenetic analysis of three gene regions (TrnL-F, rpL-16 and ITS) based on 83 sequences
were done in this study to infer the phylogeographic history of this genus. The results from
this study indicate two independent events of colonization each corresponding roughly to the
two broadly distributed species and the two narrow endemics. The morphological, phenetic
and phylogenetic analysis of this study support the current taxonomic classification of the
tropical species of the genus Pentaschistis.
vi
TABLE OF CONTENTS
DECLARATION .......................................................................................................................... II
DEDICATION ............................................................................................................................. III
ACKNOWLEDGEMENT .......................................................................................................... IV
ABSTRACT .................................................................................................................................. V
DEFINITION OF TERMS ......................................................................................................... IX
LIST OF TABLES...................................................................................................................... XIII
CHAPTER ONE: GENERAL INTRODUCTION .....................................................................1
1.1 BACKGROUND INFORMATION .......................................................................................1
1.2 EASTERN AFRICAN MOUNTAINS ...................................................................................3
1.2.1 Diversity and Distribution of flora on Eastern African Mountains ..................................7
1.2.2 Taxonomy of the genus Pentaschistis (Nees) Spach in the Tropical Eastern Africa ........8
1.3 ECONOMIC VALUE OF AFROALPINE PLANTS IN EASTERN AFRICA ................9
1.4 PROBLEM STATEMENT ....................................................................................................10
1.5 HYPOTHESES .......................................................................................................................10
1.6 OBJECTIVES OF THE STUDY ..........................................................................................11
1.6.1 General objective ..................................................................................................................11
1.6.2 Specific objectives .................................................................................................................11
1.7 JUSTIFICATION AND SIGNIFICANCE OF THE STUDY ............................................12
CHAPTER TWO: DIVERSITY AND THE DISTRIBUTION OF AFROALPINE
VASCULAR PLANT SPECIES IN EASTERN AFRICA ......................................................13
2.1 LITERATURE REVIEW ......................................................................................................13
2.1.1 Geological History ................................................................................................................13
2.1.2 Biodiversity of the Vegetation of Eastern African Mountains .........................................14
2.1.3 Afroalpine zone of Eastern Africa ......................................................................................15
2.1.4 Value of Afroalpine flora .....................................................................................................20
2.2 MATERIALS AND METHODS ...........................................................................................22
2.2.1 Sampling Design ...................................................................................................................22
2.2.2 Data Analysis ........................................................................................................................30
2.2.3 Species Diversity Analysis....................................................................................................30
2.2.4 Species Richness ...................................................................................................................32
2.2.5 Species Evenness ...................................................................................................................32
2.2.6 Pairwise comparison of intervegetation communities and intermountains using
Jaccard’s Similarity Index ..........................................................................................................32
2.2.7 Multivariate Analysis of species composition between the vegetation communities
and between mountains ...............................................................................................................34
2.2.7.1 Non−Parametric Multidimensional Scaling (MDS) ............................................ 34
2.2.7.2 Cluster Analysis ................................................................................................... 35
2.2.8 Vegetation Cover ..................................................................................................................35
2.2.9 Analysis of Variance (ANOVA) for Species richness ....................................................3636
2.2.10 Analysis of Similarities of the species composition of vegetation communities and
between mountains (ANOSIM) ..............................................................................................3636
2.2.11 Species Contribution to Similarity (SIMPER) ............................................................3636
2.3 RESULTS ..............................................................................................................................3838
2.3.1 Diversity and Distribution of Afroalpine Plants in Eastern Africa .............................3838
2.3.2 Similarities between the five vegetation types ...............................................................4040
2.3.3 Similarity between the Afroalpine species of six Mountains of Eastern Africa .........4444
vii
2.3.4 Species Diversity in the Afroalpine Vegetation communities .......................................5050
2.3.5 Comparison of Species Diversity, richness and Abundance in the Afroalpine zone of
Eastern Africa ..........................................................................................................................5454
2.3.5.1 Analysis of Similarities (ANOSIM) and Species Contribution ....................... 5858
2.3.5.2 Species Contribution to Similarity (SIMPER) among the vegetation communities
...................................................................................................................................... 6060
2.3.5.3 SIMPER among mountains.............................................................................. 7373
2.3.6 Community Description ...................................................................................................8181
2.4 DISCUSSION AND CONCLUSION ..................................................................................8585
2.4.1 Species dominance in the Afroalpine vegetation of Eastern Africa .............................8585
2.4.2 Description of the communities of the five vegetation types of Afroalpine zone ........8787
2.4.3 Comparative assessment of species composition between and within five vegetation
types of Afroalpine zone of Eastern Africa............................................................................8888
2.4.4 Comparative assessment of species richness and abundance between the five
vegetation types ........................................................................................................................9595
2.4.5 Comparative assessment of species composition between the alpine zone of the six
mountains of Eastern Africa ...................................................................................................9696
2.4.6 Heterogeneity Assessment ...........................................................................................100100
CHAPTER THREE: PHENETIC AND PHYLOGENETIC ANALYSIS OF THE
GENUS PENTASCHISTIS (NEES) SPACH (POACEAE) OCCURRING IN EASTERN
AFRICAN MOUNTAINS ......................................................................................................102102
3.1 LITERATURE REVIEW ..................................................................................................102102
3.1.1 Distribution and Description of the genus Pentaschistis ...........................................102102
3.1.2 Nomenclatural background of the genus Pentaschistis .............................................103103
3.1.3 Previous phylogenetic analysis ....................................................................................105105
3.2 MATERIALS AND METHODS .......................................................................................108108
3.2.1 Sampling design and raw data collection ...................................................................108108
3.2.2 Sampling for phylogenetic analysis.............................................................................109109
3.2.3 Phenetic analysis ...........................................................................................................109109
3.2.4 DNA analysis .................................................................................................................109109
3.2.5 Phylogenetic analysis ....................................................................................................111111
3.3 RESULTS ............................................................................................................................112112
3.3.1 Cluster analysis (CA) ...................................................................................................112112
3.3.2 Principal component analysis (PCA) ..........................................................................114114
3.3.3 Distribution and Ecological Analysis of Pentaschistis Species across the vegetation
communities ..........................................................................................................................116116
3.3.4 Molecular Analysis .......................................................................................................117117
3.4 DISCUSSION AND CONCLUSION ................................................................................120120
3.4.1 Phenetic and morphological analysis ..........................................................................120120
3.4.2 Phylogenetic analysis ....................................................................................................122122
3.4.3 Key to Pentaschistis species occurring in African high mountains .........................125125
CHAPTER FOUR: SUMMARY DISCUSSION, CONCLUSIONS AND
RECOMMENDATIONS ........................................................................................................134134
4.1 SUMMARY DISCUSSION ...............................................................................................134134
4.2 CONCLUSIONS .................................................................................................................140140
4.3
RECOMMENDATIONS .......................................................................................140140
viii
REFERENCES .........................................................................................................................141141
APPENDIX 1A: DATA MATRIX OF 12 PLOTS OF MT KENYA SHOWING
PRESENCE/ABSENCE SCORES FOR ALL SPECIES SAMPLED ...............................148148
APPENDIX 1B: DATA MATRIX OF 11 PLOTS OF MT ELGON SHOWING
PRESENCE/ABSENCE SCORES FOR ALL SPECIES SAMPLED ...............................159159
APPENDIX 1C :DATA MATRIX OF 12 PLOTS OF MT KILIMANJARO SHOWING
PRESENCE/ABSENCE SCORES FOR ALL SPECIES SAMPLED ...............................170170
APPENDIX 1D: DATA MATRIX OF 12 PLOTS OF MT RWENZORI SHOWING
PRESENCE/ABSENCE SCORES FOR ALL SPECIES SAMPLED ...............................181181
APPENDIX 1E: DATA MATRIX OF 16 PLOTS OF MT BALE SHOWING
PRESENCE/ABSENCE SCORES FOR ALL SPECIES SAMPLED ...............................192192
APPENDIX 1F: DATA MATRIX OF 12 PLOTS OF MT SIMEN SHOWING
PRESENCE/ABSENCE SCORES FOR ALL SPECIES SAMPLED ...............................204204
APPENDIX 2A: DATA MATRIX OF THE SPECIES OF PENTASCHISTIS FOR
PHENETIC ANALYSIS ........................................................................................................215215
APPENDIX 2B: DATA MATRIX ..........................................................................................217217
APPENDIX 3: SPECIES COUNT FOR PENTASCHISTIS SPECIES FROM FIVE
VEGETATION TYPES..........................................................................................................219219
APPENDIX 4: SUMMARY OF THE FAMILY, GENERA AND SPECIES SAMPLED
FROM EASTERN AFRICA ..................................................................................................220220
APPENDIX 5: COMPARISON OF SPECIES DIVERSITY & RICHNESS AMONG
THE 75 PLOTS FROM EASTERN AFRICAN MOUNTAINS.........................................222222
APPENDIX 6: FACTOR COORDINATES OF THE VARIABLES, BASED ON
CORRELATIONS (Α=0.05) SHOWING CONTRIBUTION OF EACH CHARACTER224224
APPENDIX 7: DATA MATRIX FOR SPECIES RICHNESS OF EACH VEGETATION
COMMUNITY ........................................................................................................................226226
APPENDIX 8: DATA MATRIX FOR SPECIES RICHNESS OF EACH EAST
AFRICAN MOUNTAIN ........................................................................................................227227
APPENDIX 10: PLATES OF SOME TYPICAL AFROALPINE SPECIES FROM
EASTERN AFRICA…. ..........................................................................................................235235
APPENDIX 11: BOTANICAL ILLUSTRATIONS OF PENTASCHISTIS SPECIES.....238238
APPENDIX 11: BOTANICAL ILLUSTRATIONS OF PENTASCHISTIS SPECIES......239239
ix
DEFINITION OF TERMS
Abiotic: refers to the non−living components of an ecosystem, such as water, air, soil etc.
Abundance: The number of organisms in a population that show degree of density (within
inhabited areas) and prevalence.
Acaulescent: With no stem or reduced stem
Afroalpine: Vegetation communities occurring above the treeline in high tropical African
Mountains and dominated by relatively sparse low growing vegetation with characteristic life
forms such as tussock grasses, acaulescent herbs and rosette perennials.
Afromontane: the vegetation zone below ericaceous belt dominated by tropical moist forest
in African Mountains and highlands.
Alpha diversity: The number of species recorded within some standardized area e.g. a square
kilimetre or some naturally delineated patch of habitat such as grassland, bog, a patch of
forest etc.
Anemochory: Passive dispersal of propagules by wind.
Anthropogenic: Actions or factors attributed to humans or human induced effects.
Archipelago: A group of islands often scattered in expansive water e.g. Lamu archipelago
Arctic: Pertaining to all non−forested areas north of the coniferous forests of the Northern
Hemisphere.
Beta diversity: The dissimilarity or turnover in species composition between local ecological
communities.
Biotic: pertaining to the living components of an ecosystem (from the simplest to the most
complex organisms).
Biogeography: The science that attempts to document and understand spatial and temporal
distribution of organisms. Modern biogeography now includes studies of all patterns of
geographic variation in life, from genes to entire communities and ecosystems.
x
Boreal: occurring in the temperate and sub−temperate zones of the Northern Hemisphere that
typically contains coniferous forests and some forms of deciduous forests.
Caldera: is a cauldron-like volcanic feature usually formed by the collapse of land following
a volcanic eruption. They are sometimes confused with volcanic craters. The word comes
from Spanish caldera, and this from Latin caldaria, meaning "cooking pot" (Wikipedia
encyclopedia).
Circumboreal: occurring in the temperate/sub temperate zones of the New and Old World
parts of the Northern Hemisphere.
Clade: Any monophyletic evolutionary branch in a phylogeny, using derived characters to
support genealogical relationships.
Cladogram: A line diagram derived from a cladistic analysis showing the hypothesized
branching sequence of a monophyletic taxon.
Colonization: The immigration of a species into new habitat followed by successful
establishment of a population.
Community: An assemblage of organisms that live in a particular habitat and interact with
one another.
Diaspore: Any part or stage in the life cycle of an organism that is adapted for dispersal.
Dispersal: The movement of organisms away from their point of origin.
Ecosystem: The set of biotic and abiotic components in a given environment.
Ecology: The study of the abundance and distribution of organisms and of the relationships
between organisms and their biotic and abiotic environments.
Edaphic: pertaining to soil.
Endemic: Pertaining to a taxon that is restricted to the geographic area specified e.g. a
continent, a mountain, lake, island etc.
Evolution: Any irreversible change in the genetic composition of a population.
xi
Extinct: No longer living, opposite of extant.
Gene: The small unit of a DNA molecule that codes for a specific protein to produce on of
the chemical, physiological, or structural attributes of an organism.
Habitat: Place where a microorganism, plant or animal lives.
Habitat diversity: The range of habitats present in a region.
Heterogeneity: The state of being mixed in composition, as in genetic or environmental
heterogeneity.
Immigration: Entry of organisms to a population from elsewhere.
In-group: The focal monophyletic group in a cladistic analysis.
Microclimate: The climate within a very small area or in a particular, often tightly defined,
habitat.
Migration: The movement of individuals, and commonly whole populations from one region
to another.
Miocene: A geological era lasting from approximately 25 to 5 million years ago.
Moorland is a type of habitat, in the temperate grasslands, savannas, and shrublands biome,
found in upland areas, characterised by low-growing vegetation on acidic soils and heavy
fog.
Oligocene a geological era in the middle of the Tertiary Period (and end of the Paleogene),
from approximately 33.9 to 23 million years ago.
Orogeny: The process of mountain building resulting from the upward thrust of Earth’s crust
due to volcanic or tectonic activities.
Out-group: A taxon related to the in-group, used in a cladistic analysis to infer primitive and
derived character states in a transformation series.
Pan tropical: Occurring in all major tropical areas around the world.
Phenetic: The study of the overall similarities of organisms.
xii
Phylogeny: The evolutionary relationships between an ancestor and all its known
descendants.
Phylogeography: An approach in biogeography that studies the geographic distributions of
genealogical lineages, within species and among similar species, and attempts to differentiate
between historical and ongoing processes leading to the development of observed patterns.
Pleistocene: A geological era lasting from approximately 2 million to 10 000 years ago.
Pliocene: A geological era lasting from approximately 5 to 2 million years ago.
Precambrian: The Precambrian period is that long stretch of geological time that starts from
the formation of the Earth up to the beginning of the Cambrian – about 4.6 billion years ago.
Sister group: In a phylogeny, the group or clade most closely related to the focal group (in
group), and therefore the most useful out-group for rooting the phylogeny.
Species diversity: An index of community diversity that takes into accounts both species
richness and the relative abundance of species.
Species richness: The number of species present in a community.
Stochastic: Random, expected (statistically) by chance alone.
Sclerophyllous: Having tough, thick, evergreen leaves.
Scrubland: Any of a wide variety of vegetation types dominated by low shrubs; in
exceedingly dry locations.
Taxon (plural: Taxa): A general term for any taxonomic category e.g. a species, genus,
family etc.
Vicariants: two disjunct and phylogenetically related species that are assumed to have been
created when the initial range of their ancestor was split by some historical event.
Xerophytes: Land plants that grow in relatively dry (“xeric”) environments.
xiii
LIST OF TABLES
TABLE 1: Flora elements of the Afroalpine zone of Eastern Africa .............................................19
TABLE 2: 15 Dominant families of the afroalpine plant species from Eastern Africa ...............3939
TABLE 3: Species diversity, richness, abundance and evenness in five vegetation types in six
Eastern African mountains ............................................................................................................50
TABLE 4: Post hoc tukey test indicating source of variation between some of the vegetation
communities of the Eastern Africa ............................................................................................5252
TABLE 5: Intermountain species diversity, richness, abundance and evenness .........................5555
TABLE 6: Tukey test outcome indicating source of variation between Eastern African
mountains based on the species richness ....................................................................................5757
TABLE 7: Pairwise comparison of five vegetation communities using ANOSIM .....................5858
TABLE 8: Species contribution to similarities (SIMPER) in the five vegetation types of
Afroalpine zone......... ....................................................................................................................61
TABLE 9: Pairwise comparison between the five vegetation community types from six
Eastern Africa countries ..............................................................................................................6262
TABLE 10: Summary of the differences between the six mountains of Eastern Africa … ..........71
TABLE 11:
Most important species contributing to similarities in the individual six
mountains and pairwise comparison among Eastern Africa .......................................................7474
TABLE 12: Summarry of community description from five Afroalpine vegetation
types………. ...............................................................................................................................8282
TABLE 13: A summary of the species of Pentaschistis recognized by Clayton, Phillips and
Linder in East African and Ethiopian mountains .........................................................................105
TABLE 14: Gene region amplified and sequenced and the primers used for amplification .....110110
TABLE 15: Distribution of Pentaschistis species in Eastern African mountain .......................116116
xiv
TABLE 16: Frequency of Pentaschistis species in the five vegetation communities of
Afroalpine zone of Eastern Africa ............................................................................................117117
xv
LIST OF FIGURES
FIGURE 1: Showing the five life forms in the Afroalpine zones of the high mountains ..............17
FIGURE 2A: Map of East Africa and Ethiopia showing major mountain systems and Rift
Valley........ ....................................................................................................................................23
FIGURE 2B: Afroalpine sampling points for the bog, Dendrosenecio and rock outcrop plots
from Mt. Kenya.............................................................................................................................24
FIGURE 2 C: Afroalpine sampling points for the Dendrosenecio, grassland and rock outcrop
plots from Mt. Elgon .....................................................................................................................25
FIGURE 2 D: Afroalpine sampling points for the bog, grassland and rock outcrop plots in Mt.
Kilimanjaro ...................................................................................................................................26
FIGURE 2 E: Afroalpine sampling points for Alchemilla, bog and Dendrosenecio plots of Mt.
Rwenzori ......................................................................................................................................27
FIGURE 2 F: Afroalpine sampling points for Alchemilla, bog, grassland and rock outcrop
plots of Mt. Bale ...........................................................................................................................28
FIGURE 2 G: Afoalpine sampling points of bog, grassland and rock outcrop plots of Mt.
Simen...... ......................................................................................................................................29
FIGURE 3: Percentage family contribution of Afroalpine species from Eastern Africa .............3939
FIGURE 4: Species accumulation curve showing species sampling from 75 plots ....................4040
FIGURE 5: Similarities between five vegetation community types in Eastern African
mountains ....................................................................................................................................4141
FIGURE 6: Dendrogram showing vegetation type similarity in Afroalpine zone with respect to
species composition ....................................................................................................................4343
FIGURE 7: Similarity between the high mountains of Eastern Africa…………. .......................4545
FIGURE 8: Species composition similarity between Eastern African mountains .......................4747
xvi
FIGURE 9: Multidimensional scaling graph showing species composition similarities among
Eastern African mountains ..........................................................................................................4848
FIGURE 10: Multidimensional scaling graph showing distribution of sampled plots from
Eastern African mountains ..........................................................................................................4949
FIGURE 11: Variations of the species richness of the five vegetation communities as indicated
by the vertical lines .....................................................................................................................5151
FIGURE 12: Species richness in five vegetation types in Afroalpine zone of Eastern
Africa…… ..................................................................................................................................5353
FIGURE 13: Line graph showing species abundance among the afroalpine vegetation
communities ................................................................................................................................5353
FIGURE 14: Line graph showing abundance between the six Eastern African mountains.........5555
FIGURE 15: Variations of the species richness of the six mountains of Eastern Africa as
indicated by the vertical lines ....................................................................................................5656
FIGURE 16: ANOSIM graph for the five vegetation types showing R−values ..........................5959
FIGURE 17: Comparison between species composition of the rock outcrop plots from Eastern
African mountains .......................................................................................................................6666
FIGURE 18: Comparison between species composition of the grassland plots from Eastern
African mountains .......................................................................................................................6767
FIGURE 19: Comparison between species composition of the bog plots from Eastern African
mountians ....................................................................................................................................6868
FIGURE 20: Comparison between species composition of the Dendrosenecio plots from
Eastern African mountains as revealed by MDS graph ..............................................................7070
FIGURE 21: Worldwide distribution of the species of Pentaschistis .......................................102102
FIGURE22: Phenogram of Custer analysis for Pentaschistis species .......................................113113
xvii
FIGURE 23: Scatter plot of the PCA results of morphological variations for Pentaschistis
species…. ..................................................................................................................................115115
FIGURE 24: Showing the occurrence and prevalence of five Pentaschistis species across five
vegetation types in Eastern African mountains ........................................................................117117
FIGURE 25 Phylogenetic tree inferred from Bayesian analysis of the plastid DNA and ITS
data set- posterior probability values are indicated above the branches. ..................................119119
xviii
LIST OF PLATES
Plate 1 Carduus keniensis....................................................................................................224
Plate 2 Anemone thomsonii .............................................................. ……………………235235
Plate 3 Cardamine obliqua...................................................................................................224
Plate 4 Alchemilla johnstonii ............................................................ ……………………235235
Plate5 D. advinalis var. petiolatus........................................................................................225
Plate 6 Disa stairsii .......................................................................... ……………………236236
Plate 7 Galium ruwenzoriense .............................................................................................225
Plate 8 Haplocarpha rueppellii ......................................................... …………………..236236
Plate 9 Gladiolus watsonioides ...........................................................................................226
Plate 10 Ranunculus oreophytus .........................................................…………..…….237237
Plate 11 Helichrysum formosissimum .............................................................................226
Plate 12 Hebenestreitia angolensis ......................................................……………….226
Plate 13 Galium acrophyum .............................................................................................227
Plate 14 Helichrysum forskahlii ...............................................................……………...227
Plate 15 Subularia monticola...........................................................................................227
Plate 16 Huperzia saururus ....................................................................…………….238238
1
CHAPTER ONE: GENERAL INTRODUCTION
1.1 Background Information
Any isolated habitat that is entirely separated from others of its kind can be considered as an
island. For example, there are many mountains in the world surrounded by low−lying land.
Just as oceanic islands are separated from one another by water, so mountain summits are
separated by ‘seas’ of low−lying land. Hence, mountain species have to arrive on a mountain
by some form of long−distance dispersal in much the same way as mainland species arriving
on oceanic islands. Some of the most interesting isolated “sky−islands” occur in the
mountains of the tropical regions where the upper zones are surrounded by montane forests or
savanna. The mountains of Eastern African region are like a cluster of small “sky islands”
and are mostly solitary volcanic mountains on or near the Equator surrounded by ‘sea’ of
savanna or are formed by uplift e.g. Ruwenzori (Hedberg, 1970).
The high African mountains in the equatorial zone, including the unique afro-alpine
ecosystem, act as major water towers and basically are essential for water supply and
agriculture in many African countries. These include, but are not limited to, Kenya, Uganda,
Tanzania, Burundi, Rwanda, and Ethiopia. The major mountain systems that serve as water
towers include Bale, Simen, Choke, Gara Muleta (Ethiopia), Rwenzori, Virunga cluster of
mountains (Uganda/Democratic Republic of Congo), Kilimanjaro, Meru (Tanzania),
Aberdare ranges, Mt. Kenya and Elgon (Kenya) among others. Usually the vegetation of
these mountains is divided into several distinct zones: the bamboo zone, afro-montane zone,
the sub-alpine ericaceous zone and the afro-alpine zone. In three mountains, viz: Kilimanjaro,
Kenya and Rwenzori, above the afroalpine zone occurs alpine desert and the mountains are
clad with glittering ice caps or glaciers. The afro-alpine zone typically occurs above 3500
2
metres in most of the mountains and harbours unique and beautiful alpine grasslands, shrubs,
and bogs and is home to many rare and endemic species. It has been shaped through a
dramatic history of repeated climate changes and is ideal as a natural laboratory for the study
of biodiversity dynamics. The above named mountains basically constituted the study area
and greater description of the same is given in the subsequent sections.
Generally speaking biodiversity is the variation of living organisms within a defined area
such as habitat, an ecosystem, biome, or for the entire Earth. The biodiversity occurring in it
consists of millions of both distinct and cryptic biological species, which have evolved over a
long geological period of time.
From biogeographic point of view, gradients of diversity for most organisms decrease from
the Equator towards either side of the poles. This means majority of the species of organisms
of all kind are concentrated along the Equator or generally around the tropics and
sub−tropics. Similarly diversity along altitudinal gradient increases towards mid elevation
where it reaches the peak before it gradually declines towards the mountain summit. Thus the
mountain systems including those of Eastern African high mountains mimic the globe in the
distribution patterns of plant and non− plant communities.
The importance of the surrounding biodiversity to man and other organisms is immense and
nearly impossible to quantify since it provides essential functions and services that directly
impact life on earth. Besides providing food, medicines, clothes, construction materials, dyes
etc. they play direct roles in the regulation and composition of atmospheric gases, regulation
of the global climate, generation and maintenance of soils and nutrient recycling.
3
Unfortunately anthropogenic activities including over-exploitation of species, spread of alien
species to new environments, direct habitat destruction for domestic or industrial purposes,
and pollution among others have caused huge losses of biodiversity and extinction of
numerous species. Afroalpine species are particularly vulnerable since the impact of man’s
activities has been compounded by the impact of global warming. Hence detailed study of the
Afroalpine species, their distribution, abundance, species richness and evenness will provide
baseline information necessary to give mitigation measures for the conservation of this fragile
ecosystem.
1.2 Eastern African Mountains
Mount Kenya is an extinct volcano about 3.5 million years old and straddles the Equator and
is 175 km north−east of Nairobi (Kenya Wildlife Service− KWS, 1997). It covers an area of
715 km2 and was gazetted as a National Park in 1949 transcending several administrative
boundaries including Nyeri, Kirinyaga, Embu, Meru South and Meru Central districts. Its
vegetation consists of bamboo forest, afromontane forest, sub−alpine ericaceous zone,
Afroalpine habitats and is crowned by alpine deserts and the glittering twin peaks of Batian
(5,199 m) and Nelion (5,188 m). It was declared as a World Heritage Site in 1997 for “one of
the most impressive landscapes of Eastern Africa with its rugged glacier−clad summits,
Afroalpine moorlands and diverse forests which illustrate outstanding ecological processes”
(KWS, 1997).
The Aberdare Mountain, which is a national park, is an isolated range that forms the eastern
wall of the rift valley running north to south over a distance of more than 100 km (KWS,
1997). It covers an area of 767 km2 and includes as part of the administrative boundaries of
the districts of Nyeri, Muranga, Thika, Gatundu, Laikipia West and Nyandarua. There are
two main peaks, Oldoinyo Le Satima, which is the highest peak (4,001 m), and Kinangop
4
peak (3,900), separated by a long saddle of alpine moorland that ranges over 3,000 m (KWS,
1997).
Mount Elgon is a 24 million year old extinct volcano that has eroded over time showing an
expansive caldera and is located 470 km North West of Nairobi, in the Tranzoia District of
the Rift Valley province. It is a national park that straddles the Kenya−Uganda border
covering an area of 169 km2 with altitudinal range of 2,100−4,301 m and is famous for its
bat−filled caves (KWS map, undated).
The Rwenzori (also written as Ruwenzori) Mountain lies almost at the equator, rising over
4000 m above the floor of the Albertine Rift Valley and is bisected by the Uganda−Congo
border, which passes through Mt. Stanley, the highest peak. The park trail head at
Nyakalengija is 375 km from Kampala and can be reached through Fort Portal. The
Ruwenzori Mountain is remarkable for its flora and an ascent over this mountain passes
through a series of increasingly spectacular changes of vegetation zones. Above the Bakonzo
farmlands occur montane forest (1500−2500 m) that changes to bamboo stands and tangles of
Mimulopsis (2,500−3000 m). This is followed by giant tree−heathers (3000−4000 m) starting
with Ericaceous species, and crowning with spectacular forms of giant Lobelia and
Dendrosenecio species interspersed by thickets of Helichrysum species (Uganda Wildlife
Authority− UWA Map, 2007).
Mount Muhavura is an extinct volcano in the Virunga Mountains on the border between
Rwanda and Uganda. At 4, 127 m, Muhavura is the third highest of the eight major
mountains of the mountain range, which is part of the East African Rift Valley. Muhavura
lies partly in Volcanoes National Park, Rwanda and partly in Mgahinga Gorilla National
5
Park, Uganda where parts of this study (taxonomic sampling jointly with AFROALP II
Project) was done. It is a home to the critically endangered mountain Gorilla listed on the
IUCN red list of endangered species (UWA Map, 1997)
Mount Kilimanjaro with its three volcanic cones, Kibo, Mawenzi, and Shira, is an inactive
volcano in northern Tanzania rising 4,600 m from its base covering an area of 388,500 ha.
The highest point in Africa is on Mt. Kilimanjaro and is called Uhuru peak (5,895 m), on the
volcano Kibo. Although it is inactive, Kilimanjaro has fumaroles that emit gas in the crater
on the main summit of Kibo. Mt. Kilimanjaro has unique vegetation such as the water
holding cabbage (Dendrosenecio species), Lobelia species, alpine grassland or tussocks and
other alpine vegetation all adapted in living in alpine conditions. Subalpine Erica forests at
4,100 m represent the highest altitude cloud forest in Africa. In addition it has a large variety
of forest species over an altitudinal ranges of 3,000 m containing more than 1,200 vascular
plant species (Newmark, 1991).
Mount Meru is located east of the Great Rift Valley and about 40 km southwest of
Kilimanjaro in northern Tanzania's Arusha National Park. This mountain is considered an
active volcano and is the country's second highest mountain. It is the youngest volcanic
mountain in tropical Africa and about 0.5 million years ago, Mount Meru erupted in a huge
explosion that destroyed its cone shape and resulted in a horseshoe crater (Meru crater). In
the past century, Mt. Meru had minor eruptions and the Ash Cone continues to build inside
the crater (TANAPA, 2007).
The Simen Mountain is part of the Northwestern Ethiopian Mountain systems or Highlands
and lies in the North Gondar Administrative Zone, approximately 110 km northeast of the
6
town of Gondar. It consists of broad, undulating plateau, generally above 3,000 m above sea
level and reaching over 4,000 m in some areas (Silki, Bwahit and Ras Dashan), part of which
is a National Park (Simen Mountain National Park). The tallest peak is Ras Dashan (4,453 m)
and is World Heritage site. There are two major vegetation types in the Simen Mountains.
These are Afroalpine vegetation in the uplands and Afromontane vegetation in the lowlands.
Ericaceous forest, scrub is a transitional, occurring both on the escarpments and in the
uplands (Puff & Nemomissa, 2005).
The Bale Mountains are a range of mountains in the Oromia Region of southeast Ethiopia.
They include peaks like Tullu Demtu which is the second highest point in Ethiopia (4,377 m),
and Mount Batu at an altitude of 4,307 m. These mountains are part of Bale Mountain
National Park and cover 2,200 km2. Bale Mountains contains three distinct ecoregions: the
northern plains, bush and woods; the central Sanetti Plateau (where the sampling was done)
with an average elevation of over 4,000 m that is characterized by Afroalpine vegetation
(grasslands, rocky outcrops, bogs and Alchemilla communities) and the southern Harenna
Forest famous for its mammals, amphibians and birds including many endemic species. The
central Sanetti Plateau is home to the largest population of the rare and endangered Ethiopian
red wolves. (ONRS, 2008).
Gara Muleta Mountain is an isolated, rocky mountain with an elevation of 3,405 m. It is 40
km due east of the city of Harar in the Hararghe Region of Ethiopia. The vegetation of this
Mountain helps in the conservation of the soil, water and biotic diversity. Previous vegetation
studies documented about 361 vascular plant species (Teketay, 1996). Currently the
vegetation of this mountain is increasingly being threatened by destruction caused by the
surrounding population through deforestation, fire, cultivation and overgrazing.
7
The Choke Mountains are to the south of Lake Tana in the centre of Gojam, Ethiopia. This
large mountain block rises gradually from the surrounding plateau to around 2,800 m, with
the highest peak, Mt. Choke reaching an elevation of 4,070 m. The most remarkable feature
of this mountain is the virtual absence of forest. The major natural habitats are moist
moorland with giant Lobelia species, shrubby Alchemilla species, sedges and tussocks of
Festuca species and other grasses (Birdlife International, 2010).
1.2.1 Diversity and Distribution of flora on Eastern African Mountains
Almost always vegetation changes in orderly manner as one climbs up a mountain from the
foothill to the summit of the high mountains. Eastern African Mountains are no exception and
several distinct zones of vegetation occur, viz: the savanna at the foot of the mountain
followed by the bamboo zone (in most of the mountains), and the tropical rain forest zone
occupies mid to high altitude (1500− 2800 m). This gradually gives way to the ericaceous
zone that is dominated by the genus Erica L. marking the end of the tree line zone, above
which occurs the distinctive afroalpine flora. Although this zone occurs in the tropical
equatorial zone, it is not only isolated ecologically but has a special temperate flora and is
peculiarly adapted to the extreme conditions of temperature variation.
Each mountain enclave is also isolated from seemingly neighbouring mountains and can be
viewed as archipelagos− like oceanic islands− that vary in area and isolation. Speciation rates
and endemicity increase with both area and isolation hence the very high endemicity of
Afroalpine flora (above 80%) Hedberg (1970). The main plant communities occurring in
these mountains are: Dendrosenecio forests/woodland, tussock grassland, Carex bogs and
8
sclerophyllous scrub (Helichrysum/ Alchemilla scrub). Because of the high isolation of the
Afroalpine zone, relatively few collections have been made and the area is seldom visited by
researchers as it is both inhospitable and expensive to do botanical research in this zone. As
such there is little or no known detailed account and comparative study of how diversity of
plants species, species richness, abundance and evenness vary between the various plant
communities of each mountain and between mountains. The study will also boost the herbaria
collections of many taxa, which either are altogether lacking or are severely under collected.
1.2.2 Taxonomy of the genus Pentaschistis (Nees) Spach in the Tropical Eastern Africa
Pentaschistis (Nees) Spach consists of about 70 species, of which the majorities are
indigenous to Cape Floristic Region (CFR) and other parts of South Africa. The genus is an
African endemic and belongs to the tribe Danthonieae of the sub-family Danthonioideae
(Poaceae). Both the generic and species delimitation of this genus have been variously
described as difficult by different authors (Chippendall, 1955; Clayton & Renvoize, 1986;
Linder & Ellis, 1990). Consequently the various species of Pentaschistis, particularly those
occurring in South Africa, were placed in different genera over the years, such as in Holcus
L., Avena L., Danthonia DC., Eriachne R. Brown., Triraphis R. Br. and Achneria Munro ex
Benth. & Hook. f. Similarly the species circumscription of the tropical African Pentaschistis
varied over the last 75 years and this therefore necessitated taxonomic revision. This is due
to the insufficient collections and high degree of variability in the tropical African species. As
a result the population on each of the various high mountain enclaves was initially treated as
different species by Hubbard (1937). He recognised four species among the Pentaschistis
with open panicles: P. ruwenzoriensis C.E. Hubb. from Ruwenzori, P. meruensis C.E. Hubb.
from Mt. Meru, P. expansa (Pilg.) C.E. Hubb. from Mt. Kenya and P. borussica (K. Schum.)
Pilg. from several mountains. However all these were later united under the name P.
9
borussica by Clayton (FTEA, 1970). Phillips (1994, 1995) reduced P. minor, P. mannii, P.
gracilis to variety levels under P. pictigluma.
Hence this study used various methods such as phenetic and phylogenetic techniques in
addition to morphology both using herbarium specimens and field observations of different
populations to carry out taxonomic revision of this genus to address the question of differing
species circumscription.
1.3 Economic value of Afroalpine plants in Eastern Africa
Economic botany can be defined as the exploitation of plants and plant products by humans
for domestic or commercial purposes. Traditional societies including both pastoral and
farming communities, particularly in Africa and Asian countries, have in-depth knowledge of
the plant biological resources surrounding their environment (Timberlake, 1987) and
frequently utilize them for both domestic and commercial purposes. The most commonly
known uses of plants include: source of foods such as cereals, nuts, edible fruits and
vegetables; source of medicine, construction materials such as timber and bamboo; fibres
such as cotton for textile industry; dyes such as tannins; source of poisons such as African
arrow poison; fuel such as firewood and charcoal; shade and as ornamentals among others.
Many researchers in Eastern Africa have documented traditional plant names and uses.
Kokwaro (1972, 1976) has compiled numerous plant names and uses from all Kenya and East
Africa. Glover et al. (1966; 1969) have also documented Maasai and Digo plant names and
uses. The most common and significant use of plant resources is as a source of traditional
medicines to treat many ailments and conditions. Many pharmaceutical drugs globally used
today like artemisinin-based antimalarial drugs currently used in Kenya (from genus
10
Artemisia), thymol in Listerine (mouth wash) from the genus Thymus and atropine from the
genus Atropa among many others have come from folk use and use of plants by indigenous
cultures. As a result, numerous ethnobotanical studies aimed at identifying new
pharmaceutical products have been initiated recently (Balick, 1994).
Due to the extreme isolation of Afroalpine zones of the Eastern Africa, coupled with the fact
that all the mountains are protected as national parks, few of the Afroalpine plants are used
for various purposes compared to other plant species from forest or lowland ecosystems by
the surrounding communities. However quite a number of the Afroalpine species have uses
ranging from source of medicine to cultural uses.
1.4 Problem statement
The study involves assessment of species richness, species diversity and abundance in the
five vegetation types viz. rock outcrop, bog, grassland, Dendrosenecio forest/woodland and
Alchemilla communities of six Eastern African mountains: Mt Kenya, Kilimanjaro, Elgon,
Rwenzori, Bale and Simen. In addition the study compares the similarities of species
composition between the five vegetation types and mountains as well as addressing the
species circumscription of the genus Pentaschistis in the Afroalpine zone of Eastern Africa.
1.5 Hypotheses
•
There are no significant differences in the plant species richness between rock
outcrop, bog, grassland, Dendrosenecio and Alchemilla vegetation communities
•
There are no significant differences in the plant species richness between Mt. Kenya,
Kilimanjaro, Elgon, Ruwenzori, Bale and Simen
•
There are no significant differences in terms of plant species composition between the
six mountains.
11
•
There are no significant differences in the five vegetation types (communities) of
Afroalpine zone of Eastern Africa
•
Phenetically and morphologically there are no significant differences between the
variations of all the population of Pentaschistis in the mountains of Eastern Africa
•
There is only single colonization event of the genus Pentaschistis in tropical African
Mountains
1.6 Objectives of the study
1.6.1 General objective
Determine the diversity and distribution of the Afroalpine species in Eastern Africa as well
evaluate the species circumscription of the tropical species of the genus Pentaschistis
1.6.2 Specific objectives
•
Compare vascular plant species diversity between the various vegetation communities
of the Afroalpine zone of each of the six mountains (alpha and beta diversity)
•
Compare the similarity of plant species composition between the various vegetation
communities of each mountain and between mountain systems
•
Determine the species richness, abundance and evenness of Afroalpine plant species
in the various vegetation communities
•
Test heterogeneity (distinctness) of vegetation communities
•
Establish the diagnostic features of Tropical African Pentaschistis and determine how
many species should be recognized
•
Evaluate the phenetic relationship between the species of Pentaschistis in the study
area
12
•
Determine the distribution and frequency of Pentaschistis species in the five
vegetation communities of Afroalpine zone of Eastern Africa
•
Revise the phylogeny of the Pentaschistis clade of Danthonioideae based on plastid
and nuclear DNA sequences.
1.7 Justification and significance of the study
There is no detailed research work on the floristic composition, species diversity analysis,
abundance, species richness and similarity and the conservation status of the main vegetation
types of the Afroalpine zone of these mountains. Since the scope of these mountain enclaves
constituting the Afroalpine zone is both limited in area and isolated detailed information on
its species is urgently needed for conservation purposes because of not only high endemicity
exhibited by its species but also the rarity of many Afroalpine species as well as the fragility
of this peculiar ecosystem.
The genus and species of Pentaschistis has undergone various shifts within Danthonioideae
and hence has been referred to as a difficult (Chippindal 1955; Clayton & Renvoize, 1986;
Hilliard & Burtt, 1987; Phillips, 1994). This has been aggravated by the insufficient
collections and high degree of variability in the tropical African species, and the huge
geographical separation between populations in the mountain refugia. Therefore species
circumscription has been a problem. Hence this study was done to assess the diversity and
distribution of the vascular plants of Afroalpine species in six mountain systems of East
Africa and Ethiopia as well as revise the taxonomy of the genus Pentaschistis. It will provide
a comprehensive species checklist that will be useful in the conservation and management of
this fragile ecosystem.
13
CHAPTER TWO: DIVERSITY AND THE DISTRIBUTION OF AFROALPINE
VASCULAR PLANT SPECIES IN EASTERN AFRICA
2.1 LITERATURE REVIEW
2.1.1 Geological History
The present day landscape in Africa owes its origin to a combination of geological processes
mainly operating in opposition. On the one hand, continuous weathering, erosion and
deposition of the Mozambique belt have resulted in leveling of the land thereby producing
extensive plains in much of the African continent (Clifford, 1970). On the other hand
sporadic faulting has elevated some portions and lowered others. In some areas this faulting
has been accompanied by volcanism (Clifford, 1970). From the Precambrian (>600 million
years ago) until the onset of rifting during the Miocene, most part of the African continent
was little affected by major earth movement (Hamilton, 1982). The middle tertiary
(approximately 25 million years ago) witnessed the beginning of the doming of extensive
areas of Central and Eastern Africa (Gautier, 1967; Morgan, 1973).
These geological events produced extensive plains in the eastern half of Africa sometimes
stretching for hundreds of kilometres. Above these plains occur isolated hills called
inselbergs as well as very high mountains, of which, three reach above the height where
plants cannot survive (Mt. Kilimanjaro, Mt. Kenya and Mt. Ruwenzori). These mountains
and others in this region resulted from the Recent Pleistocene−Pliocene rift tectonics and
volcanism, approximately between 0.01 and 7 million years ago (Clifford, 1970).
Consequently, there is a wide range of landscape features in Eastern Africa including
Ethiopia as a result of these complex geological processes associated with intensive volcanic
activity and faulting that gave rise to the Great Rift Valley (Nyamweru, 1980). The rift is a
14
giant crack running through the African continent starting from the Afar depression in
northern Ethiopia to almost at the mouth of river Zambezi in Mozambique covering a
distance of 5,600 km. The accompanying volcanic activities are responsible for the creation
of many mountains that are distributed along the faults of this great valley. This has
profoundly changed the physiographic appearance of the region such that some spectacular
scenery is found in countries such as Ethiopia, Kenya, Uganda, Tanzania, Burundi and
Rwanda (Hedberg, 1970).
2.1.2 Biodiversity of the Vegetation of Eastern African Mountains
Biological diversity or simply biodiversity includes all species of plants, animals and
microorganisms including their genetic material as well as the ecosystems or habitats in
which they occur. It is the central axis to the study of biology (Darwin, 1872; Howard &
Berlocher, 1998; Dickson & Foster, 2011) and in particular it is the core topic of ecological
theory and conservation biology (Pärtel et al., 2010). Biodiversity directly and indirectly
affects human development (Lusweti, 2011) and is therefore crucial for the socio−economic
welfare of mankind. Human induced global climate change is causing ecological
communities to lose species (Wardle et al., 2011) through accelerated rate of extinction
especially fragile ecosystems such as Afroalpine vegetation types in mountain system. It is
therefore paramount to thoroughly study species diversity, abundance, cover and distribution
to pinpoint the rare, endangered and vulnerable species so that appropriate measures can be
taken to conserve them. In addition this can influence policy makers towards the conservation
of these vulnerable components of biodiversity. Therefore currently, biodiversity
conservationists are pre−occupied with species loss from ecosystems caused by habitat
destruction mostly related to anthropogenic factors (Wilson & Peter, 1988; Stuart et al., 1990;
Glowka et al., 1994; IUCN, 1994; UNEP, 2000a). Hence information on the distribution and
15
abundance of biological species especially the rare and endangered ones are of primary
importance in the planning and implementation of biodiversity conservation (Condit et al.,
1996; Eilu et al., 2004b).
Communities including those of Afroalpine flora have distinctive ecological and physical
boundaries as shown by the composition, height and growth form among others.
Communities and related natural phenomena such as soil, climate and topography too,
continuously co−vary (Mueller−Dombois & Ellenberg, 1974; Whittaker, 1975; Ritter, 2001).
While tropical forests are well known for being the most species−rich ecosystems on earth
(Gentry, 1992) Afroalpine ecosystems are thought to be species poor as is evident from the
phytochorion in which they are described (“…Afroalpine archipelago−like region of extreme
floristic impoverishment” (White, 1978a)).
Most of the previous studies on vegetation ecology of Afroalpine flora in Eastern Africa are
based on the work of Olov Hedberg (Hedberg, 1951, 1955, 1957, 1961, 1964 and 1970) but
other recent works also exist and include: accounts on Mt. Kilimanjaro (Beck et al., 1983),
Mt. Kenya (Rehder et al., 1981 & 1988; Beck et al., 1990; Bussmann, 1994; Bussmann &
Beck, 1995a), Aberdare range (Schimitt, 1991), Mt. Ruwenzori (Schimitt, 1992), Mt. Elgon
(Wesche, 2002), Virunga cluster of mountains (Fischer & Hinkel, 1992), Mt. Bale (Gashaw
& Fetene, 1996; Miehe & Miehe, 1993, 1994b; Bussmann, 1997), Mt. Simen (Sileshi & Puff,
2005).
2.1.3 Afroalpine zone of Eastern Africa
The vegetation of the high Mountains of East Africa and Ethiopia displays conspicuous
zonation, starting with a montane forest belt, followed by an ericaceous belt and an
16
Afroalpine belt (Hedberg, 1951). The Afroalpine region was originally proposed by Hauman
(1955) to comprise the Afroalpine calottes of the equatorial Eastern Africa, notably the
Virunga volcanoes, Ruwenzori, Elgon, Mt. Kenya, Kilimanjaro and Mt. Meru. Later on
Ethiopian high mountains like Simen and Bale were added to the Afroalpine regions. Despite
the close proximity, the flora of the Afro−alpine is floristically distinct and ecologically
isolated from the surrounding montane forest and savanna vegetation. Consequently it has
been described by Hedberg as isolated temperate “Sky–islands” since the low lying montane
and savanna vegetation separate the mountain enclaves from each other. The Afro−alpine
parts of these mountain systems of eastern Africa are characterized by large variations of
temperature during the day and night, that is, “summer every day and winter every night”
(Hedberg, 1970), very high degree of endemism of flora and fauna; are species poor
compared to the surrounding montane and savanna vegetation communities. The Afro−alpine
zone has four major vegetation types: Bogs, rock outcrop, alpine grassland/moorland and
mix scrubland (Alchemilla community or Dendrosenecio forests/woodland) and its plant
community have been categorized into five distinct life forms adapted to tropical alpine
conditions: giant rosette plants (giant Lobelia and Dendrosenecio species), tussock grasses,
acaulescent rosette plants (Haplocarpha species), cushion forming plants (Subularia
monticola) and sclerophyllous plants (Erica and Alchemilla species, Hedberg, 10) (Figure 1).
17
Figure 1: Showing the five life forms in the Afroalpine zones of the high mountains: a) giant rosette plant; b) Tussock grass; c)
acaulescent rosette plant; d) cushion forming plants; and e) Sclerophyllous plants (Source: Hedberg, 1964b).
18
The Afroalpine flora is very interesting because of the striking ecological adaptations
exhibited by many species (Hedberg, 1964) and its large amount of geographically vicarious
taxa. Its giant Dendrosenecio and giant Lobelias are as renowned in this respect as the finches
of the Galapagos Islands. Some 80% of the Afroalpine species of vascular plants are endemic
to the high mountains of East Africa and Ethiopia (Hedberg, 1961). Vicarious taxa occur of
different status. Sometimes one species occurs on all or most of the Eastern Africa with
vicariad in other parts of the world e.g. Subularia monticola A. Br. ex Schweinf. (Afroalpine)
and S. aquatic L. (Circumboreal). In other cases each of two species is confined to one group
of mountains, as in the species pair Lobelia wollastonii E. G. Bak. (Virunga volcanoes and
Ruwenzori) and L. telekii Schweinf. (Elgon, Aberdare, Mt. Kenya). Finally, there are several
groups of vicarious taxa where each taxon is endemic or limited to one mountain, as in
Lobelia deckenii (Asch.) Hemsl. group with six cognate species. The vicarious species of the
Afroalpine flora have arisen as the result of geographical isolation. The massive mountains
harbouring them are of unequal ages (Miocene to late Pleistocene) and have stood isolated
from each other since their origin (Hedberg, 1961). Their Afroalpine enclaves are therefore
markedly isolated from each other. Phytogeographically, species of Afroalpine plants are
categorized into nine flora elements based on the supposed areas of origin of each species,
that is, the areas in which they have the majority of their closest relatives (Hedberg, 1961).
These floral elements are summarized in table 1.
19
Table 1: Flora elements of the Afro alpine zone of Eastern Africa
Group
No. of Taxa
Endemic Afroalpine
52
element
Endemic afromontane
35
element
South African element
17
Cape element
10
South Hemispheric
11
temperate element
North Hemispheric
43
temperate element
Mediterranean element
18
Himalayan element
5
Pantemperate element
87
Source: (Hedberg, 1961)
Percentage (%)
19
13
6
4
4
15
6
2
31
The endemic Afroalpine element consists of monotypic or oligotypic genera that are entirely
or almost entirely confined to the Afroalpine belt e.g. Keniochloa Melderis, Oreophyton O. E.
Schulz, Haplosciadium Hochst. and Nannoseris Hedb. among others. Also included within
this category are members of such genera that have their main distribution in the Afroalpine
belt like Dendrosenecio and Alchemilla L. species. The endemic afromontane element
consists of members of genera whose distribution mainly occurs at lower levels (afromontane
zone) such as giant Lobelia L. species. The third group, the South African element contains
species that have their closest relatives in South Africa outside the Cape region e.g. Dierama
pendulum (L.f.) Bak. The Cape element category consists of Afroalpine species that have
their closest relatives in the Cape region proper e.g. Stoebe kilimandscharica O. Hoffm. and
Pentaschistis (Nees) Spach species. The fifth group, the South Hemispheric temperate
element includes members whose relatives occur in the Antarctic region e.g. Nothofagus .
The North Hemispheric, or Boreal element comprises 43 taxa whose relatives are
concentrated in Europe e.g. Cardamine hirsuta L., Arabis alpina L, Callitriche stagnalis
20
Scop. among others. Members of the genera of the Mediterranean element have their relatives
in the Mediterranean region and include such species as Erica arborea L., Aira caryophyllea
L., Romulea fischeri Pax, Trifolium multinerve (Hochst.) A. Rich., Anthemis tigreensis J. Gay
ex A. Rich. etc. The smallest element in the Afroalpine flora is the Himalayan element,
consisting of taxa that have their nearest relatives in the mountains of Southern Asia. It
contains only five taxa: Satureja punctata (Bth.) Briq., S. biflora (Buch.−Ham. ex D. Don)
Briq., Dicrocephala alpina R. E. Fr., Crepis scaposa R. E. Fr. ssp. afromontana (R. E. Fr.)
Babc., C. suffruticosa Babc. The ninth and the largest group is the Pan temperate element and
comprises species that either occur themselves on several continents in both hemispheres, and
that have their closest relatives so distributed. Examples in this category include
Deschampsia flexuosa (L.) Trin., Festuca kilimanjarica Hedb., F. pilgeri St.–Yves ssp.
pilgeri, F. abyssinica Hochst. ex A. Rich. etc (Hedberg 1961).
2.1.4 Value of Afroalpine flora
Plants are a major component of the biodiversity and are essential in maintaining life on our
planet since they manufacture food in the form of organic compounds such as starch, fats and
vitamins, among others, which are utilized by man and other animals for sustenance
(Neuwinger, 1996). Consequently man is known to have been using wild plant resources
since time immemorial to satisfy a wide range of basic needs of both rural and urban
communities, in both tropical and sub-tropical countries (UNESCO report, 1998) including
those living around the high mountains of Eastern Africa. Besides being a source of
sustenance, plants also produce numerous other products, usually called secondary
metabolites, that were used since ancient times for both domestic (mostly traditional
medicine) and commercial purposes (for example essential oils for perfume manufacture and
21
ingredients of pharmaceutical products). Thus the field of economic botany is one of the
oldest disciplines in botanical science.
Traditional medicine in Kenya and other parts of Eastern Africa has been practiced for many
centuries and still plays an important role in providing primary health care (Kokwaro, 1988;
1976) due to the inadequate health facilities and few trained health personnel available. This
is evident by the fact that about 75% of the Kenyan population for example relied on
traditional medicine as their primary source of health care and over 90% of the people have at
one time or another resorted to the use of herbal medicine (Odera, 1997). It is for this reason
that the World Health Organization (WHO) has been encouraging the use of traditional
medicine, particularly in the developing countries through promoting the incorporation of its
useful elements into national health care systems (Akerele, 1987).
According to WHO, up to 80% of the population in Africa uses traditional medicine for their
primary health care (Hamilton, 2008). The majority of plant species used in herbal medicine
treatments are harvested from the wild rather than cultivated. Therefore overexploitation
especially for commercial purposes may endanger some species, particularly those endemics
that have limited distribution. Herbal remedies may be prepared in various forms, as
infusions, decoctions, powdery ash and instillations (Lindsay, 1978). Apart from providing
spectacular scenery for tourist attraction, which boosts tourist industry, as well as playing
important ecological roles, a number of the species of the Afroalpine vegetation in the
Eastern Africa are used for purposes such as source of medicine for humans and livestock,
firewood, food or beverage preservation and cultural/ritual activities among others.
22
2.2 MATERIALS AND METHODS
2.2.1 Sampling Design
Four different vegetation community types were sampled and analysed in each of the 11
mountains viz.: Simen, Choke, Bale, Gara Muleta (Ethiopia), Kenya, Aberdare, Elgon
(Kenya), Meru, Kilimanjaro (Tanzania), Ruwenzori and Muhavura (Uganda) (figures 2 a–
2g).
These
were
rock
outcrops,
bogs,
grassland,
and
mixed
scrubland
Dendrosenecio/Alchemilla). In each vegetation type in each mountain, four plots of
approximately 100×100 m were sampled and analysed. Raw data (herbarium voucher
specimens and silica dried leaves) were collected from the plots using the Plot and No−plot
protocols developed by the Afroalp II project (NUFU project 2007/1058).
For each population of plant species, five plants were sampled, one from each of the four
corners of the plot and one from the middle, that is, complete sampling of plant species was
done. Leaves from these plants were put in tubes containing silica gel and the first three
plants were taken as vouchers for morphological reference and verification. In case the
species were not occurring within the plot, a sample of five individuals of each species was
collected from outside the plots to represent the mountain (s). Plant identification was done
both in the field and in the herbarium using available local floras (Flora Tropical East Africa
and Flora of Ethiopia and Eritrea) as well as relevant botanical books (Beentje, 1994;
Agnew & Agnew, 1994) in addition to expertise of participating taxonomists. Most of the
collections were identified to species level and variety/sub−species level where applicable.
23
Figure 2a: Map of East Africa and Ethiopia showing major mountain systems and Rift Valley,
which formed the study area (Source: AFROALP II Project)
24
Figure 2b: Afroalpine sampling points for the bog, Dendrosenecio and rock outcrop plots from Mt. Kenya
25
Figure 2 c: Afroalpine sampling points for the Dendrosenecio, grassland and rock outcrop plots from Mt. Elgon
26
Figure 2 d: Afroalpine sampling points for the bog, grassland and rock outcrop plots in Mt. Kilimanjaro
27
Figure 2 e: Afroalpine sampling points for Alchemilla, bog and Dendrosenecio plots of Mt. Rwenzori
28
Figure 2 f: Afroalpine sampling points for Alchemilla, bog, grassland and rock outcrop plots of Mt. Bale
29
Figure 2g: Afoalpine sampling points of bog, grassland and rock outcrop plots of Mt. Simen
30
2.2.2 Data Analysis
The completeness of the sampling (species checklist) was assessed using the accumulative
species curve computed using PRIMER version 5.0 (Clarke & Warwick, 1994; Clarke &
Gorley, 2001).
The software calculates species sampling data matrix by iteratively
resampling the raw data 999 times and averaging the result.
2.2.3 Species Diversity Analysis
Whittaker (1972) classified diversity into alpha (α), beta (β) and gamma (γ) types. The α
diversity is that of species within habitat(s) or community, β diversity is the measure of the
rate and extent of change in a species along a gradient from one habitat to others while γ
diversity is the richness in species of a range of habitats in a geographical area. Many
diversity indices have been proposed (example reviews by Eberhardt, 1969; Peet, 1974;
Magurran, 1988; Krebs, 1999; Southwood & Henderson, 2000 and Feinsinger, 2001) and
there is no consensus on which one of them is the best (Lewis et al., 1988; Krebs, 1999).
Diversity measures are divided into species richness measure, species abundance and indices
based on the proportional abundance of species (Magurran, 1988). Peet (1974) suggests that a
number of concepts are lumped within the term diversity e.g. species richness, evenness (or
equitability) and heterogeneity (combines species richness and evenness). There are two
types of heterogeneity indices commonly referred to as type 1 and type 2 (Peet, 1974). Type
1 is the most sensitive to changes in the importance of rare species in the sample (example is
Shannon−Wiener index) while type 2 are the most sensitive to changes in the most abundant
species (example Simpson index). Evenness indices include Pielou’s (J’) index, Lloyd and
Ghelardi’s index (ε) [Lloyd & Ghelardi, 1964; Magurran, 1988). Importance of diversity
indices include: −
31
1. Illustrates intrinsic diversity patterns of ecological communities and Ecosystems
2. Indicates the relative contribution of abundant and rare species to diversity
3. Shows successional trends of a community over time as affected by abundant and rare
species (Lewis et al., 1988)
However diversity indices have limitations that can affect the accuracy of a monitoring
protocol (Norris, 1999) because of their sensitivity to sample size and bias, taxonomic
uncertainties and stochastic sampling.
This study used Shannon−Wiener diversity index (H’) computed to base e using PRIMER
version 5.0 (Clarke & Warwick, 1994; Clarke & Gorley, 2001) of the data matrix of the
species sampling from the 75 plots (Appendix 1a– 1f).
H’= nΣpi (log2) pi,
i=1
Where n is the number of species, pi is the proportion of the total count emanating from the
ith species (Clarke & Warwick, 1994).
The index is computed on the basis of what
proportion of the total individuals of each species comprises. Advantages of this index are: −
•
Useful in describing ecological trends because it includes both species count and
evenness
•
Have moderate discriminant ability
•
An intermediate ease of calculation
Since it is difficult to interpret diversity indices, it is recommended to combine with other
measures of diversity in giving correct interpretation and description of ecological
communities. These include measures such as total number of species (species richness),
abundance and evenness.
32
2.2.4 Species Richness
Species richness was quantified by employing Margalef’s index (d) [Clifford & Stephenson,
1975) using the formula:
d= (S−1)/log N where S is the total number of species and N is the total number of
individuals. The advantage of using this index is that it minimises the effect of sample sizes
and the fact that S and N are simple and easy to compute. The data matrix used was the
present/absent scores for all the species sampled from the 75 plots.
2.2.5 Species Evenness
In this study, Pielou’s evenness index (J’) was used (Pielou, 1975). It shows how evenly the
individuals occurring are distributed among the different species. The data matrix used was
the present/absent scores for all the species sampled from the 75 plots. The index ranges
between 0 and 1 with 1 representing even distribution of individuals within all the species
while lower values (towards 0) indicate dominance by one or few individual taxa (uneven
distribution). The index (J’) is calculated as: J’= H’ (observed)/H’max where H’max is the
maximum possible diversity achievable when all species are equally abundant (Pielou, 1975;
Clarke & Warwick, 1994). Its advantage includes the fact that it reduces dependence on
sample size and simplicity of its computation.
2.2.6 Pairwise comparison of intervegetation communities and intermountains using
Jaccard’s Similarity Index
Jaccard’s Similarity Index was used to compare the degree of similarity between the five
vegetation community types in the Afroalpine Mountains of Eastern Africa (Jaccard, 1901).
Each time two vegetation communities were compared and the values of the common
(shared) species, different and Jaccard’s computed using Microsoft excel data analysis sheet.
33
The pooled data matrix of the scores of all species present/absent in each vegetation
community from the 75 plots was used. It was computed from the equation: j/ (a+b-j)
Where: ‘j’ is the number of species found in both samples and ‘a’ is the number of species in
sample a while b is the number of species in sample b.The higher the value of Jaccard’s the
more similar two vegetation communities are (in terms of species composition).
A total of 10 comparisons are possible between the five vegetation community types (rock
outcrop Vs. bog, rock outcrop Vs. grassland, rock outcrop Vs. Dendrosenecio, rock outcrop
Vs. Alchemilla, bog Vs. grassland, bog Vs. Dendrosenecio, bog Vs. Alchemilla, grassland
Vs. Dendrosenecio, grassland Vs. Alchemilla and Dendrosenecio Vs. Alchemilla). Similarly
the species composition among the six mountains (Mt. Kenya, Elgon, Kilimanjaro,
Ruwenzori, Bale and Simen) was compared using Jaccard’s similarity index. It resulted with
15 pairwise comparisons (Mt. Kenya Vs. Elgon, Kenya Vs. Kilimanjaro, Kenya Vs.
Ruwenzori, Kenya Vs. Bale, Kenya Vs. Simen, Elgon Vs. Kilimanjaro, Elgon Vs.
Ruwenzori, Elgon Vs. Bale, Elgon Vs. Simen, Kilimanjaro Vs. Ruwenzori, Kilimanjaro Vs.
Bale, Kilimanjaro Vs. Simen, Ruwenzori Vs. Bale, Ruwenzori Vs. Simen and Bale Vs.
Simen.
34
2.2.7 Multivariate Analysis of species composition between the vegetation communities
and between mountains
2.2.7.1 Non−Parametric Multidimensional Scaling (MDS)
In order to do comparisons in the species composition of all the five vegetation communities
and the six mountains at the same time, non−parametric Multidimensional scaling (MDS)
was used. MDS gives a good link between the original data and the final picture and
represents complex patterns correctly in low dimensional space (Clarke & Warwick, 1994). It
only considers that an ordination is important representation of similarity by looking at the
stress values that range from 0 to 1. The lower the stress value the better the two dimensional
presentation of the data point and ideally values 0.1 or less gives the best presentation.
Similarity matrix was first generated which was used to produce graphs to discriminate sites
from each other and cluster sites with similar species composition (Clarke & Warwick,
1994). In this study, ordinations were calculated using Primer version 5.0 where the original
data matrix was first converted to Bray−Curtis matrix using the SIMPER module. This is the
most commonly used similarity coefficient used in ecological analysis. It neutralises the
effect of the rare species (common in Afroalpine flora) such that the rare a species is, the less
it contributes to the overall pattern (Clarke & Warwick, 1994). Comparisons of species
composition using MDS was done between the 75 plots, five vegetation communities and the
six mountains. The aggregation of the various points (plots, vegetation communities or
mountains) depends on the degree of similarity of species composition of the points.
35
2.2.7.2 Cluster Analysis
Cluster analysis of the same data (from plots, vegetation types and mountains) were
performed to compare with the results of those from MDS since similar plots or vegetation
types or mountains are expected to cluster together and give easily interpretable dendographs.
The Bray−Curtis Algorithm was used to generate the data matrix and dendographs (Clarke &
Warwick, 1994).
2.2.8 Vegetation Cover
Ecological assessment for each plot in the five sub−plots were also done that involved
estimates of total vegetation cover, cover of vascular plants, bryophytes, lichens and
individual species of vascular plants (percentage cover) and information recorded in the Plot
protocol. In the eleven mountain systems in which raw data (percentage cover of each species
in the sub–plots) was collected, it was only possible to collect plot data from six mountains
viz: Mt. Bale, Simen (Ethiopia), Mt. Kenya, Elgon (Kenya side), Mt. Kilimanjaro (Tanzania)
and Mt. Ruwenzori (Uganda) and therefore a total of 75 plots were sampled and data
obtained for ecological assessment. The cover data was useful in the community description
observed during the sampling of the plots from all the sampled mountains. Only taxonomic
sampling was done in the remaining five mountains:
Choke Gara Muleta (Ethiopia),
Aberdares (Kenya), Mt. Meru (Tanzania) and Muhavura/Gahinga (Uganda). Hence the
analysis of the data in this section only covers data obtained from plots. A reference
collection of voucher specimens from each species sampled (in all the plots and non−plot
samples) are deposited at EA herbarium (Kenya), national herbarium in Addis Ababa
(Ethiopia), Makerere University herbarium (Uganda) , national herbarium in Arusha
(Tanzania) and Oslo herbarium (Natural History Museum, University of Oslo, Norway).
36
2.2.9 Analysis of Variance (ANOVA) for Species richness
The data sets from vegetation communities and mountains were used to discern whether there
were significant variations between them using ANOVA. First the pooled data matrices of
species richness in each plot in the respective vegetation communities and mountains
(Appendix 7 and 8 respectively) were rearranged to suit the STATISTICA program then
Margalef’s species richness (d) from the five vegetation communities and mountains was
subjected to single factor (one−way) ANOVA using STATISTICA version 7 (Statsoft Inc.,
1999). Tukey’s multiple comparison tests were performed to find the source of variations
whenever ANOVA tests were significant (Tukey, 1977).
2.2.10 Analysis of Similarities of the species composition of vegetation communities and
between mountains (ANOSIM)
ANOSIM test allow a statistical test (1−way layout) of the null hypothesis that there are no
assemblage differences between groups of samples, specified a priori (Clarke & Gorley,
2001). The key output is in the results log and a histogram of ANOSIM R statistic values
which are obtained from 999 random permutations of the groups (mountains or vegetation
communities in this study for example) between the samples and shows the likely values of
R. These are distributed around zero−if there are no differences between groups than
between−group and within group similarities will be roughly equal−but they are never more
than 0.15 or so by chance. The data matrixes of the species present/absent of the 75 plots
were used to compare assemblage differences between the vegetation communities and
between mountains (Heterogeneity test).
2.2.11 Species Contribution to Similarity (SIMPER)
It is possible to identify which species primarily account for the observed assemblage
differences of the various vegetation types (and mountains). This can be done by the
37
decomposition of Bray−Curtis similarity (for within treatments e.g. within plots of any given
mountain or vegetation community) (or dissimilarity) of between treatments (e.g. between 2
or more mountains or vegetation communities) index into contribution from each species.
That is, by looking at the overall percentage contribution of each species makes to the
average dissimilarity between two groups. Hence one list species in decreasing order of
importance in discriminating the two sets of samples. This can be done using SIMPER,
particularly Bray−Curtis similarity matrix (Clarke & Gorley, 2001). Here no standardization
and Transformation is done and a cut−off percentage of 90% is chosen. This means that
species will be listed in decreasing order of their importance, in contributing to the average
dissimilarity between two groups, until the point is reached at which 90% of the dissimilarity
is accounted for (Clarke & Gorley, 2001). The data matrixes of the present/absent species
from the 75 plots were used to assess the contribution of each species to the overall
similarities in the vegetation communities and mountains.
38
2.3 RESULTS
2.3.1 Diversity and Distribution of Afroalpine Plants in Eastern Africa
A total of 278 species from 124 genera belonging to 46 families were sampled from 75 plots
in five vegetation community types (Appendix 4 and 9) viz: rock outcrop, bogs, grassland,
Dendrosenecio forest/woodland and Alchemilla community from the four mountains of East
Africa (Mt. Kenya, Elgon, Kilimanjaro and Ruwenzori) and two Ethiopian Mountains (Bale
and Simen). About 58 specimens were not fully identified to species level and are excluded
from this number of species in the checklist (Appendix 7). However much of the known
Afroalpine species were covered in this list of (46 out of the 48 families/124 genera out of
141 genera).
Of the 46 families collected from the plots 15 families (Asteraceae, Poaceae,
Caryophyllaceae, Apiaceae, Brassicaceae, Scrophulariaceae, Crassulaceae, Cyperaceae,
Iridaceae, Adiantaceae, Ranunculaceae, Rubiaceae, Fabaceae, Gentianaceae and Lobeliaceae)
constituted 81% of all the species recorded (figure 3) i.e. out of the 278 species, they
constitute 180 species (Table 2) (Appendix 2). Asteraceae represented by 21 genera out of
124 genera(17.36%) and 69 species out of 278 species (24.82%) and Poaceae with 17 genera
(14.05%) and 37 species [13.31%]) were the most dominant taxa of the Afroalpine vegetation
communities in the six mountains sampled followed by Caryophyllaceae (9 genera−7.44%;
16
species−5.76%),
Apiaceae
(7
genera−5.79%;
9
species−3.24%),
Brassicaceae
(5genera−4.13%; 6 species−2.16%), Scrophulariaceae (5 genera−4.13%; 8 species−2.88%),
Crassulaceae (5 genera−4.13%; 8 species−2.88%), Cyperaceae (4 genera−3.31%; 11
species−3.96%), Ranunculaceae (3 genera−2.48%; 9 species−3.24%), Adiantaceae (3
genera−2.48%; 4 species−1.44%) and Iridaceae (3 genera−2.48%; 3 species−1.08%).
39
Table 2: 15 dominant families of the Afroalpine plant species from Eastern Africa
No. of Percentage No. of Percentage
No. Family
genera (%)
Species (%)
T
47
124
see below
278
See below
1
Asteraceae
21
17
69
25
2
Poaceae
17
14
37
13
3
Caryophyllaceae
9
7
16
6
4
Lamiaceae
4
3
12
4
5
Cyperaceae
4
3
11
4
6
Rosaceae
2
2
10
4
7
Ranunculaceae
3
2
9
3
8
Lobeliaceae
1
1
9
3
9
Apiaceae
7
6
9
3
10 Scrophulariaceae
5
4
8
3
11 Gentianaceae
1
1
8
3
12 Fabaceae
2
2
8
3
13 Crassulaceae
5
4
8
3
14 Brassicaceae
5
4
6
2
15 Rubiaceae
1
1
4
1
Figure 3: Percentage family contribution of Afroalpine species from Eastern Africa
40
Enough sampling was achieved as revealed by the average species accumulation curve for the
75 sampled plots. Here the shape of the species curve indicates the expected pattern of
initial exponential increase (sharp rise) in species with increasing sampling, then gradually
slowing down (as demonstrated by the curvature) as more samples are added until the curve
attains almost flat curvature where more sampling hardly adds new previously unaccounted
for species (Figure 4).
(
)
Sp e cie s C o u n t
(C u mu la tive )
300
250
200
150
100
50
0
1 3 5 7 9 111315171921232527293133353739414345474951535557596163656769717375
2 4 6 8 101214161820222426283032343638404244464850525456586062646668707274
Number of Plots (Samples)
Figure 4: Species Accumulation Curve showing species sampling from 75 plots
2.3.2 Similarities between the five vegetation types
Jaccard’s Similarity Index between the five vegetation community types in the Afroalpine
zone of Eastern Africa showed that rock outcrop and grassland were the most similar in terms
of species composition with Jaccard’s similarity index of 51.98. Similarly bogs and grassland
showed relatively high similarity index (44.03 Jaccards) while the most dissimilar vegetation
communities were those of between grassland Vs. Dendrosenecio vegetation communities
(21.62 Jaccards), rock outcrop verse Alchemilla (26.19 Jaccards) and rock outcrop Vs.
Dendrosenecio vegetation communities (27.36 Jaccards) (figure 5).
41
Figure 5: Similarities between five vegetation community types in Eastern Africa mountains
42
Multivariate analysis of the five vegetation community types using Cluster analysis
(Bray−Curtis) exhibits similar patterns with that of pairwise comparison (Jaccard’s similarity
index). Rock outcrop, bog and grassland cluster together (Figure 6).
43
60
Alchemilla
Dendrosenecio
Grassland
100
Rock
80
Bog
Bray Curtis Similarity index
40
Vegetation Communty
Figure 6: Dendrogram showing vegetation type similarity in Afroalpine zone with respect to species composition
44
2.3.3 Similarity between the Afroalpine species of six Mountains of Eastern Africa
Similarity between the mountains showed that the East Africa (Kenya, Uganda and Tanzania)
Mountains are more similar in terms of species composition compared with the Ethiopian
Mountains (Figure 7 & 8). Similarly the Ethiopian Mountains indicate similar trend where
they share more species. Among the East Africa mountains, Mt. Kenya, Kilimanjaro and
Elgon are more similar while Ruwenzori appears to separate from the rest. The former two
are the most similar as revealed by the relatively high Jaccard’s similarity index (44.4).
Similarly Bale and Simen show high degree of similarity (47.05). The most dissimilar
mountains are Simen and Ruwenzori (13.22) and Bale and Rwenzori (14.93).
45
Figure 7: Similarity between the high mountains of Eastern Africa
46
The cluster analysis like the pairwise comparison reveals that Simen and Bale have closer
affinities in terms of species composition thus clustering together (Figure 8). On the other
hand, Mt. Kenya, Kilimanjaro and Elgon form a cluster indicating higher degree of similarity.
Mount Ruwenzori, the only non−volcanic (Block Mountain) mountain occupies somewhat
intermediate position between the Ethiopian Mountains and the remaining East Africa
Mountains. The scatter plot graph from MDS exhibit same pattern of aggregation where the
mountains Kenya, Tanzania and Uganda aggregate closely compared to the other mountains
(Bale and Simen) of the sub–region of Ethiopia (Figure 9 and 10).
47
40
60
Mountain
Figure 8: Species composition similarity between Eastern Africa mountains
KILIMANJARO
KENYA
ELGON
RWENZORI
100
SIMEN
80
BALE
Bray Curtis Similarity
20
48
Stress: 0.01
Kenya
Elgon
Kilimanjaro
Rwenzori
Bale
Simen
Figure 9: Multidimensional scaling graph showing species composition similarities among Eastern Africa mountains
49
Stress: 0.21
Kenya
Elgon
Kilimanjaro
Rwenzori
Bale
Kenya,
Kilimanjaro
& Elgon
Simen
Bale & Simen
Rwenzori
Figure 10: Multidimensional Scaling graph showing distribution of sampled plots from Eastern Africa mountains
50
2.3.4 Species Diversity in the Afroalpine Vegetation communities
Shannon Wiener diversity index indicate that rock outcrop had the highest species diversity
(4.91 H’) among the five vegetation community types followed by grassland (4.79 H’) and
bogs (4.68 H’) while the Dendrosenecio vegetation community was the least species diverse
in the Afroalpine zone of the East Africa and Ethiopian Mountains (Table 3). Similarly rock
outcrop was the most species rich among the five vegetation communities (188 species)
followed by bog and grassland communities with each 157 species. Alchemilla (77 species)
and Dendrosenecio (68 species) vegetation communities were most species poor (Figure 9).
Abundance too takes similar patterns with rock outcrops (660 individuals) and bogs (586
individuals) taking the lead (Figure 10).
Comparative analysis of the species richness between the five vegetation types indicates
significant variation in terms of species count (richness) between the plots of the vegetation
communities Dendrosenecio forest/woodland (68 species) vs. rock outcrop (188 species) and
Dendrosenecio (68 species) Vs. bogs (157 species) (p < 0.05; F (4, 70) =6.0771; df = 70.0).
Similarly there is a significant variation of species richness between rock outcrop and
Alchemilla vegetation communities (Figure 11, 12, and table 4).
Table 3: Species diversity, richness, abundance and evenness in five vegetation types in
six Eastern African mountains
Sample
Rock
Bog
Grassland
Dendrosene
cio
Alchemilla
Species
richness
(S)
188
157
157
68
Abundance Margalef’
(N)
s index
(d)
660
28.8
586
24.48
436
25.67
205
12.59
Pielou’s
evennes
s (J’)
0.9381
0.9263
0.9485
0.9509
87.74
70.27
87.99
35.58
Shannon Simpson
(H’)
(1−lamb
da
4.912
0.992
4.684
0.9893
4.796
0.9919
4.012
0.9833
77
160
0.971
58.35
4.218
14.97
Fisher
0.9898
51
38
36
34
32
Species Count
30
28
26
24
22
20
18
16
14
RO C
BO G
DEN
G RA
ALC
Habit at
Figure 11: Variations of the species richness of the five vegetation communities as indicated by the vertical lines (Mean Standard error)
(p=0.0003))
52
The post hoc test for the above analysis is shown in table 4 below.
Table 4: Post hoc Tukey test indicating source of variation between some of the
vegetation communities of the Eastern African mountains based on the species richness
(Between MSE = 84.593, df = 70.000 )
Habitat
Rock
Bog
Dendrosenecio Grassland Alchemilla
0.796897
0.000796*
0.346359
0.010268*
0.014012*
0.926656
0.094797
0.116521
0.996332
outcrop
Rock outcrop
Bog
0.796897
Dendrosenecio
0.000796*
0.014012*
Grassland
0.346359
0.926656
0.116521
Alchemilla
0.010268*
0.094797
0.996332
0.370358
0.370358
The table indicates the pairs of vegetation communities that have significant variations in
terms of species richness (p< 0.05). The numbers with asterix show the p-values of the pairs
of vegetation communities with significant variations (p< 0.05) and includes rock vs.
Dendrosenecio, rock vs. Alchemilla and bog vs. Dendrosenecio (Figure 12).
53
Figure 12: Species richness in five vegetation types in Afroalpine Zone of Eastern
Africa
Figure 13: Line graph showing species abundance among the Afroalpine vegetation
communities
54
Plant species abundance similarly exhibited similar trend with species richness where rock
outcrop, bog and grassland vegetation communities respectively showing the highest
abundance. Alchemilla community showed the least abundance (figure 13).
2.3.5 Comparison of Species Diversity, richness and Abundance in the Afroalpine zone
of Eastern Africa
Among the six mountains analysed for species diversity, Simen (4.83 H’) and Bale (4.47 H’)
[Ethiopian Mountains] had the highest diversity as revealed by the Shannon Wiener’s
diversity index (parenthesis) compared to other East Africa Mountains (Mt. Kenya,
Kilimanjaro and Elgon). Mount Elgon (4.31 H’) has the highest species diversity among the
four East Africa Mountains followed by Mt. Kilimanjaro (4.12 H’) and Mt. Kenya (4.06 H’)
while Mt. Ruwenzori, the only wet mountain among the six was the least species diverse
(3.77 H’). The same trend was observed in terms of species richness where Simen was the
most species rich among the six mountains (156 species) followed by Bale (119 species) and
Elgon (93 species). Mount Ruwenzori was the most species poor (57 species) (Table 5;
Figure 15). Abundance too exhibited similar trend with Simen (515 individuals) and
Ruwenzori (239 individuals) having the most and least abundance level respectively (Figure
14 and table 5).
55
Figure 14: Line graph showing Abundance between the six Eastern African mountains
One way ANOVA (single factor) analysis between the six mountains of East Africa and
Ethiopia indicate (table 5) that species count (species richness) for Mt. Simen (156 species) is
significantly higher than the rest of the mountains (Figure 15), that is P< 0.05 (F (5, 69)
=15.794; df= df = 69.0) for all other mountains and Mt. Simen while there is no significant
differences between the species richness of other mountains (P>0.05). The post hoc test for
the above analysis is shown in table 6.
Table 5: Intermountain Species Diversity, richness, Abundance and evenness
Mountain
Kenya
Elgon
Kilimanjaro
Rwenzori
Bale
Simen
S
75
93
81
57
119
156
N
280
304
285
239
421
515
d
13.13
16.09
14.15
10.23
19.53
24.82
J'
0.94
0.95
0.94
0.93
0.94
0.96
H'
4.06
4.30
4.12
3.76
4.49
4.8
1−Lambda'
0.98
0.99
0.98
0.98
0.99
0.99
56
50
45
Species Count
40
35
30
25
20
15
10
KEN
ELG
KI L
RWE
BAL
SI M
Figure 15: Variations of the species richness of the six mountains of Eastern Africa as indicated by the vertical lines (Mean Standard
error) (F (5, 69) =15.794, p=.0.001; df= 69.00)
57
Table 6: Tukey test outcome indicating source of variation between Eastern African Mountains based on the species richness
( F[5, 69]=15.794; df = 69.000; p= 0.001)
MOUNTAIN
Kenya
Kenya
Elgon
Kilimanjaro Ruwenzori Mt. Bale
0.724574
0.999993
0.863016
0.894466
0.000129
0.801219
0.132988
0.997352
0.000146
0.795506
0.941730
0.000129
0.216009
0.000129
Elgon
0.724574
Kilimanjaro
0.999993
0.801219
Rwenzori
0.863016
0.132988
0.795506
Bale
0.894466
0.997352
0.941730
0.216009
Simen
0.000129
0.000146
0.000129
0.000129
Simen
0.000129
0.000129
The p− values in bold format indicate significant variations. Here it reveals that the species count or richness in Simen Mountain is significantly
higher than all other mountains. However the species richness between any other pair of mountain is not significantly different from each other
(p>0.05).
58
2.3.5.1 Analysis of Similarities (ANOSIM) and Species Contribution
Analysis of similarities (ANOSIM) was performed on Primer to compare the five vegetation
types (rock outcrop, bog, grassland, Dendrosenecio and Alchemilla). The automatically
generated table summarizing the computations and Global test summary statistic is given in
table 7 below.
Global Test
Sample statistic (Global R): 0.386
Significance level of sample statistic: 0.1%
Number of permutations: 999 (Random sample from a large number)
Number of permuted statistics greater than or equal to Global R: 0
Pairwise Tests
Table 7: Pairwise comparison of five vegetation communities using ANOSIM (R: 0.386;
significance level: 0.1%; No. of permutations 999)
Groups
Rock
outcrop and
Bog
Rock
outcrop and
Grassland
Rock
outcrop and
Dendrosene
cio
Rock
outcrop and
Alchemilla
Bog
and
Grassland
Bog
and
Dendrosene
cio
R
Statistic
0.469
Significanc Possible
e level (%) permutations
0.1
Too many
Actual
Number
Permutations observed
999
0
0.204
0.3
Too many
999
2
0.369
0.1
84672315
999
0
0.418
0.1
3108105
999
0
0.43
0.1
Too many
999
0
0.509
0.1
84672315
999
0
59
Bog
and
Alchemilla
Grassland
and
Dendrosene
cio
Grassland
and
Alchemilla
Dendrosene
cio
and
Alchemilla
0.394
0.1
3108105
999
0
0.474
0.1
13037895
999
0
0.259
0.9
735471
999
8
0.257
1.8
75582
999
17
The pair wise tests comparisons of ANOSIM assessment between the five vegetation types
reveal significant difference between the pairs (vegetation communities are recognizable or
are heterogeneous), with significance level less than 5% in all the vegetation types (0.1 to 1.8
%) (Table 7). However the R values which gives an absolute measure of how separated the
groups are (Figure16), on a scale of 0 (indistinguishable) to 1 (all similarities within
vegetation types are less than any similarities between vegetation types) show various levels
of separation.
267
247
Fre q u e n c y
181
148
59
51
1
25
9
9
1
1
-0.10 -0.08 -0.06 -0.04 -0.02 0.00 0.02 0.04 0.06 0.08 0.10 0.12 0.14
R
Figure 16: ANOSIM graph for the five vegetation types showing R−Values
60
2.3.5.2 Species Contribution to Similarity (SIMPER) among the vegetation communities
In order to compare the various vegetation communities and mountains in their species
composition, it is important to identify which species primarily account for the above
observed assemblage difference between the five vegetation types and six mountains.
Species Contribution to Similarity (SIMPER) was used to quantify the contribution of each
species in terms of percentage. The top five species from each vegetation type or mountain is
listed in a table form as well as the first ten species contributing to dissimilarity between each
pair of vegetation types or mountains. The results of the SIMPER analysis between
vegetation communities revealing the contribution of each species is given in the form of
percentage similarity/dissimilarity within and between vegetation communities in tabulation
format (Table 8 and Table 9). The first five species (contributing most to the observed
pattern) are listed for each vegetation community followed by the first ten species between
each pair of vegetation communities.
Generally the vegetation communities from East Africa Mountains (Mt Kenya, Kilimanjaro
and Mt Elgon) have closer affinities to each other as they cluster together (have more
common species). This was also true for the plots of the various vegetation types within the
Ethiopian Mountains. When species composition of single vegetation community was
compared, such as the plots of rock outcrops or bogs or grasslands, the distribution of the
plots generally takes a similar pattern where those plots from East Africa sub–region
aggregates together due to higher degree of similarities between their species (Figure 17−19).
61
Table 8: Species Contribution to Similarities (SIMPER) in the five vegetation types of
Afroalpine Zone
Species
Average
abundance
Average
similarity
Similarity/SD
Contribution
%
Cumulative
%
Helichrysum
citrispinum
Arabis alpina
Cerastium octandrum
Sagina afroalpina
Asplenium
aethiopicum
Ranunculus
oreophytus
Crassula granvikii
Cardamine obliqua
Carex monostachya
Haplocarpha
rueppellii
Koeleria capensis
Luzula abyssinica
Cerastium octandrum
Helichrysum forskahlii
Erica arborea
Cerastium octandrum
Alchemilla
argyrophylla
Galium ruwenzoriense
Festuca abyssinica
Arabis alpina
Cardamine obliqua
Peucedanum kerstenii
Helichrysum
stuhlmannii
Galium ruwenzoriense
Alchemilla
argyrophylla
ssp.
argyrophylla
0.8
1.93
1.22
6.94
6.94
0.75
0.75
0.75
0.65
1.72
1.57
1.57
1.20
1.07
1.06
1.07
0.79
6.19
5.66
5.64
4.32
13.13
18.79
24.43
28.74
0.90
2.83
1.81
8.80
8.80
0.85
0.75
0.75
0.70
2.49
1.95
1.88
1.62
1.44
1.03
1.06
0.92
7.74
6.05
5.84
5.04
16.54
22.59
28.43
33.46
0.81
0.69
0.69
0.56
0.50
0.82
0.82
2.65
1.77
1.70
0.99
0.88
3.63
3.34
1.25
0.87
0.86
0.63
0.53
1.32
1.35
11.34
7.65
7.35
4.26
3.80
10.63
9.8
11.34
18.98
26.34
30.59
34.40
10.63
20.43
0.73
0.73
0.64
0.63
0.50
0.50
2.95
2.84
2.36
1.73
1.11
1.11
0.98
0.98
0.77
0.72
0.51
0.51
8.67
8.34
6.92
7.57
4.86
4.86
29.10
37.43
44.36
7.57
12.43
17.29
0.50
0.50
1.11
1.11
0.51
0.51
4.86
4.86
22.15
17.01
62
Table 9: Pairwise Comparison between the five vegetation community types from Six Eastern Africa countries
Group rock outcrop & bog; average dissimilarity 81.64%; total no. of species 158
Average
Dissimilarity/Sta Contribution %
Species
Average Average
abundan abundance dissimilarity ndard deviation
2
ce 1
0.10
0.90
1.36
1.87
1.67
Ranunculus
oreophytus
0.00
0.75
1`.24
1.58
1.51
Carex
monostachya
0.75
0.05
1.22
1.49
1.50
Arabis alpina
0.80
0.15
1.19
1.42
1.50
Helichrysum
citrispinum
0.85
1.18
1.34
1.45
Crassula granvikii 0.25
0.00
0.70
1.14
1.40
1.39
Limosella major
Cardamine
0.15
0.75
1.14
1.31
1.39
obliqua
0.65
0.05
1.04
1.22
1.27
Asplenium
aethiopicum
0.05
0.60
1.02
1.12
1.25
Subularia
monticola
0.60
0.25
0.93
1.03
1.14
Crepis
dianthoseris
Group: Rock outcrop & grassland; average dissimilarity 79.19%; total of 150 species
0.75
0.06
1.26
1.47
1.59
Arabis alpina
Koeleria capensis
0.3
0.81
1.11
1.19
1.40
0.80
0.31
1.08
1.17
1.36
Helichrysum
citrispinum
Cumulative %
1.67
3.18
4.68
6.13
7.58
8.98
10.37
11.64
12.88
14.02
1.59
3.00
4.36
63
0.65
0.06
1.07
1.20
1.35
Asplenium
aethiopicum
0.31
1.03
1.15
1.30
Sagina afroalpina 0.75
0.60
0.44
0.91
0.96
1.15
Crepis
dianthoseris
0.45
0.50
0.88
0.94
1.11
Erica arborea
0.55
0.56
0.88
0.99
1.11
Helichrysum
forskahlii
0.55
0.44
0.87
0.97
1.09
Alchemilla
microbetula
0.40
0.44
0.87
0.94
1.09
Pentaschistis
borussica
Group: Bog & grassland; average dissimilarity 82.41%; total of 145 species
0.06
1.51
1.76
1.83
Crassula granvikii 0.85
0.90
0.25
1.35
1.40
1.64
Ranunculus
oreophytus
0.75
0.06
1.33
1.42
1.62
Cardamine
obliqua
0.75
0.13
1.28
1.37
1.55
Carex
monostachya
Limosella major
0.70
0.00
1.26
1.38
1.53
0.70
0.13
1.18
1.26
1.44
Haplocarpha
rueppellii
0.60
0.00
1.15
1.14
1.40
Subularia
monticola
0.40
0.81
1.11
1.05
1.34
Koeleria capensis
0.31
1.00
1.02
1.22
Sagina afroalpina 0.60
0.45
0.56
0.94
0.96
1.14
Helichrysum
forskahlii
Group: Rock outcrop & Dendrosenecio; average dissimilarity 79.64%; total of 127 species
0.80
0.00
1.58
1.78
1.99
Helichrysum
5.71
7.01
8.16
9.27
10.37
11.47
12.56
1.83
3.470.75
5.09
6.64
8.17
9.60
11.0
12.35
13.56
14.70
1.99
64
citrispinum
0.73
1.25
1.18
1.57
Festuca abyssinica 0.25
0.64
1.24
1.14
1.55
Cardamine oblique 0.15
0.27
1.19
1.20
1.50
Sagina afroalpina 0.75
0.30
0.73
1.18
1.14
1.48
Galium
ruwenzoriense
0.65
0.18
1.17
1.13
1.47
Asplenium
aethiopicum
0.60
0.18
1.14
1.10
1.43
Luzula abyssinica
0.40
0.82
1.10
1.07
1.39
Alchemilla
argyrophylla
0.60
0.27
1.10
1.03
1.38
Crepis
dianthoseris
0.55
0.36
1.07
0.98
1.34
Helichrysum
forskahlii
Group: Bog & Dendrosenecio; average dissimilarity 79.89%; total of 110 species
0.09
1.68
1.71
2.11
Crassula granvikii 0.85
0.90
0.18
1.64
1.60
2.06
Ranunculus
oreophytus
0.75
0.00
1.58
1.60
1.98
Carex
monostachya
0.15
0.73
1.48
1.28
1.85
Galium
ruwenzoriense
0.70
0.00
1.46
1.40
1.83
Limosella major
0.05
0.64
1.44
1.20
1.80
Arabis alpina
0.70
0.09
1.41
1.33
1.76
Haplocarpha
rueppellii
0.60
0.00
1.35
1.15
1.68
Subularia
monticola
0.73
1.27
1.07
1.59
Festuca abyssinica 0.35
Luzula abyssinica
0.65
0.18
1.26
1.16
1.58
3.56
5.11
6.61
8.09
9.56
10.99
12.37
13.75
15.09`
2.11
4.17
6.15
8.00
9.82
11.63
13.39
15.07
16.66
18.24
65
Group: Grassland & Dendrosenecio; average dissimilarity 84.82%; total of 123 species
0.81
0.18
1.65
1.40
1.95
Koeleria capensis
0.13
0.82
1.64
1.53
1.94
Alchemilla
argyrophylla
0.06
0.64
1.51
1.19
1.78
Arabis alpina
Cardamine
0.06
0.64
1.46
1.19
1.72
obliqua
0.25
0.73
1.44
1.17
1.70
Galium
ruwenzoriense
0.69
0.18
1.43
1.19
1.69
Luzula abyssinica
0.73
1.36
1.11
1.61
Festuca abyssinica 0.31
0.13
0.55
1.17
1.02
1.38
Anthoxanthum
nivale
0.55
1.16
1.00
1.37
Carduus keniensis 0.25
0.56
0.36
1.15
0.99
1.35
Helichrysum
forskahlii
Group: Rock outcrop & Alchemilla; average dissimilarity 84.04%; total of 134 species
0.80
0.25
1.28
1.28
1.52
Helichrysum
citrispinum
0.75
0.25
1.21
1.23
1.44
Arabis alpina
Sagina afroalpina 0.75
0.25
1.19
1.24
1.41
0.60
0.00
1.16
1.14
1.38
Crepis
dianthoseris
0.15
0.63
1.14
1.13
1.36
Cardamine
obliqua
0.55
0.25
1.08
1.01
1.28
H. forskahlii
0.65
0.38
1.05
1.03
1.25
Asplenium
aethiopicum
0.60
0.38
1.04
1.01
1.23
Luzula abyssinica
0.00
0.05
0.99
0.96
1.18
H. stuhlmannii
0.05
0.50
0.99
0.96
1.18
P. kerstenii
1.95
3.89
5.66
7.39
9.09
10.77
12.38
13.77
15.14
16.49
1.52
2.96
4.37
5.75
7.1
8.38
9.63
10.86
12.04
13.22
66
Stress: 0.15
Kenya
Elgon
Kilimanjaro
Bale
Simen
Figure 17: Comparison between species composition of the rock outcrop plots from Eastern Africa mountains using MDS graph
67
Stress: 0.13
Elgon
Kilimanjaro
Bale
Simen
Figure 18: Comparison between species composition of the grassland plots from Eastern Africa mountains as revealed by MDS graph
68
Stress: 0.13
Kenya
Rwenzori
Kilimanjaro
Bale
Simen
Figure 19: Comparison between species composition of the bog plots from Eastern Africa mountains as revealed by MDS graph
69
The genus Dendrosenecio was absent from Ethiopian Mountains since it is endemic to East
Africa mountains. Comparisons of the plots of Dendrosenecio show clear distinction between
Mt Ruwenzori, Elgon and Kenya. The differences between the mountains of Eastern Africa
are listed in table 10.
70
Stress: 0.13
Kenya
Elgon
Rwenzori
Figure 20: Comparison between species composition of the Dendrosenecio plots from East Africa Mountains as revealed by MDS graph
71
Table 10: Summary of the differences between the six mountains of East Africa and Ethiopia
Mountain
Nature
of
Formati
on
Age
(MY BP)
Location
Rock type
Seasonality of Mean
rainfall
Annual
rainfall
Soil type
Kenya
Volcanic
2−3
Eastern arm
Bimodal−March to
June (long rains)
and
October−December
(Short rains
1500−2500 mm
−Lithosols
characterised
by
shallow depth & stony occurring
mainly in rock outcrops
−Rankers & Gleysols: dark &
very rich in organic matter, on top
of brown, sandy, mottled loams
occur
on
valley
bottoms
(Dendrosenecio vegetation type
−Regosols−on
ridges
and
Morraines (Dendrosenecio).
Elgon
Volcanic
22
Between
eastern
and
Western(away
from the rift)
Chief rock present in
addition to Kenyte and
phonolite is basalt−are 2
varieties: −
Massive olivine−basalt of
the alpine zone (often
columnar.
Fissile
olivine−basalt−occurs in the
forest zone. Lavas of Kenya
discharged
in
order:
phonolite−Kenyte−Basalt
(igneous rocks)
Lava
of
Elgon
is
alkaline−Mainly nephilites
and phonolites. The alpine
region is covered by
agglomerates and relatively
young lava (1 % of entire
elgon rock)
Bimodal−March to
June (long rains)
and
October−December
(Short rains
890−1525 mm
Kilimanjaro
Volcanic
1−2
Eastern arm
Bimodal−March to
May (long rains)
and
250−500 mm in
the Afroalpine
Common soils are: dystric (or
Fibric) histosols with soil PH
4.5−5.7;
Poorly
developed
rankers and Regosols (profile
from 4190 was classified as
rankers but was transition to
Gleysols of lower slopes. True
bogs/peat soils with high organic
content common on valley
bottoms (bog vegetation type
Soils in Caldera are Regosols and
humic Gleysols
-Lithosols
−Rankers & Gleysols ( in valley
bottoms)
Kilimanjaro rocks include
Kenyte (foot of Kibo) &
phonolitic
trachytes,
72
Ruwenzori
Uplift
2
Western arm
Bale
Uplift/domi
ng
Miocene
and
Oligocene
(25−38)
Eastern arm
Simen
Uplift/domi
ng
25
Western arm
argitic−trachytes that often
contain
olivine,
nepheline−phonolite
and
nephelinites. (range from
sub−acidic rocks−trachytes
& phonolitic obsidians to
ultra−basic limburgite
Is a block of Eozoic rocks
(gneisis
and
schists)−
metamorphic
rocks:
Mica−schist, amphibolites,
Calc−schists,
quartzites,
Muscovites
and
Cordierite−schists
Basaltic and trachitic parent
rock, agglomerates & tuffs.
Vast quantities of basaltic
lava covered large portion
of underlying Mesozoic
rock
Trappean Basaltic lava
covered ancient Mesozoic
rock
−Regosols−on ridges and slopes
October−December
(Short rains
Rains throughout
the year (300 days)
5000 mm
Acidic
soils-Histosol
occasionally
Regosols
Rankers
and
and
Unimodal−Single
wet
season:
March−October;
November−Februar
y is dry season
800−1000
mm−alpine
zone
Andosols, Lithosols
−Rankers & Gleysols −Regosols
Unimodal: (April)
May−October
(November);
December−March
dry
season.
E−facing
slopes
receive more rain
than
W−facing
slopes
500−1500 mm
Andosols
73
2.3.5.3 SIMPER among mountains
Species contributing to the similarity/dissimilarity within and between the six mountains are
listed in table 11. The species listed in the table for each mountain and pair of mountains is
the taxa that contribute most in the observed patterns. For example in Mt. Kenya
Dendrosenecio keniodendron, Carduus schimperi, Sagina afroalpina, Lobelia telekii and
Anthoxanthum nivale are the most abundant species that frequently occur in this mountain
hence contributing greatly to the observed similarities among its vegetation communities.
Similarly, the species contributing most to the observed similarities among the vegetation
types of the Afroalpine zone of Mt. Elgon are: Galium ruwenzoriense (7.6%), Helichrysum
formosissimum (7.5%), Luzula abyssinica (5.9%), Koeleria capensis (4.93) and Geranium
arabicum (3.9%).
The average similarity between the vegetation types of Mt Kilimanjaro is approximately
40.0% and the similarity pattern is mainly contributed by the most abundant and frequently
occurring species such as Erica arborea (8.8%), Helichrysum newii (6.8%), H. forskahlii
(6.8), Luzula abyssinica (5.8%) and Euryops dacrydioides (5.6%). For Mt Rwenzori the top
five species frequently occurring in the vegetation communities are Dendrosenecio advinalis
(10.4%), Helichrysum stuhlmannii (8.4%), Alchemilla argyrophylla ssp. argyrophylloides
(8.2%) and Galium ruwenzoriense (6.7%). The average similarity between the vegetation
communities of Mt Ruwenzori is 51.6%.
In the case of Ethiopian Mountains, Bale and Simen, Colpodium hedbergii (5.2– 5.6%) and
Cerastium octandrum (4.5–5.0%) are among the species contributing to the observed
similarities between the vegetation communities of these mountains. Dicrocephala
chrysanthemifolia (6.7%), Haplocarpha rueppellii (5.6%) and Galium acrophyum (5.1%) are
74
the other frequently occurring species in the vegetation communities of Bale Mountain.
Similarly Sagina afroalpina (4.5%), Lobelia rhynchopetalum (3.8%) and Crassula granvikii
(3.5%) are the common species in Simen Mountains that contribute to the similarities of its
vegetation communities.
For the pairwise comparison between the mountain pair, Mt. Kenya and Elgon show total
dissimilarity of 74.0% that are attributed to endemic species of each mountain as well as the
frequent species in each mountain. For example D. keniodendron, which is Mt Kenya
endemic and D. elgonensis that is Mt Elgon endemic are the two most important species that
distinguishes between the vegetation communities of the Afroalpine zones of the two
mountains. Other species that account for the dissimilarity between Mt Kenya and Elgon are
the ones that are very abundant/frequent in one mountain but rare in the other mountain. For
example Carduus schimperi with average abundance of 1% and 0.2% in Mt Kenya and Elgon
respectively, Koeleria capensis with average abundance of 1% in Mt Elgon and 0.1% in Mt
Kenya , H. formosissimum with average abundance of 0.3% in Mt Kenya compared to Mt
Elgon with 1% average abundance. Other species with differential average abundance among
Mt Kenya and Elgon are: Geranium arabicum (0.1% in Mt Kenya; 0.6% in Elgon), Galium
acrophyum (0.7% in Kenya; 0.2% in Elgon), Alchemilla microbetula (0.7% in Kenya; 0.1%
in Elgon), G. ruwenzoriense (0.3% in Kenya; 0.1% in Elgon) and Sagina afroalpina (1% in
Kenya; 0.4% in Elgon) (Table 11).
Table 11: Most important species contributing to Similarities in the individual six
Mountains and pairwise comparison among Eastern Africa
Group: Mt Kenya; average similarity is 42.38%; a total of 30 species from 12 plots
Species
Average
Average
abundance similarit
y
Similarity/
Std Dev.
Contribu Cumulative %
tion %
75
1.00
4.44
5.52
10.47
10.47
Dendrosenecio
keniodendron
3.70
2.02
8.74
19.21
Carduus schimperi 0.92
0.83
2.86
1.39
6.74
25.96
Sagina afroalpina
0.75
2.35
1.05
5.55
31.50
Lobelia telekii
Anthoxanthum
0.75
2.24
1.05
5.29
36.79
nivale
Group: Mt Elgon; average similarity is 38.87%’ total of 34 species
0.91
2.94
2.01
7.56
7.56
Galium
ruwenzoriense
0.91
2.91
2.02
7.49
15.05
Helichrysum
formosissimum
0.82
2.29
1.33
5.89
20.94
Luzula abyssinica
0.73
1.92
0.99
4.93
25.87
Koeleria capensis
0.64
1.51
0.77
3.88
29.75
Geranium
arabicum
Group: Mt Kilimanjaro; average similarity 39.95%; total of 29 species
0.92
3.50
2.02
8.76
8.76
Erica arborea
0.83
2.72
1.40
6.81
15.57
Helichrysum
newii
Helichrysum
0.83
2.72
1.40
6.81
22.38
forskahlii
2.30
1.03
5.75
28.14
Luzula abyssinica 0.75
0.75
2.23
1.06
5.58
33.72
Euryops
dacrydioides
Group: Bale; average similarity 31.44%; total of 38 species
0.75
2.09
1.05
6.65
6.65
Dicrocephala
chrysanthemifolia
0.69
1.76
0.88
5.59
12.24
Haplocarpha
rueppellii
Colpodium
0.69
1.65
0.89
5.23
17.47
hedbergii
0.63
1.61
0.76
5.12
22.60
Galium
acrophyum
0.69
1.58
0.90
5.01
27.61
Cerastium
octandrum
Group: Simen; average similarity 35.62%; total of 61 species
1.00
2.35
9.00
6.61
6.61
Colpodium
hedbergii
0.92
1.97
2.14
5.52
12.13
Cerastium
octandrum
1.59
1.42
4.47
16.60
Sagina afroalpina 0.83
0.75
1.36
1.08
3.83
20.43
Lobelia
rhynchopetalum
0.75
1.26
1.07
3.53
23.95
Crassula
granvikii
Group: Ruwenzori; average similarity 51.60%; total of 18 species
1.00
5.37
3.90
10.40
10.40
Cardamine
obliqua
76
0.92
4.35
1.88
8.44
18.84
Dendrosenecio
advinalis
0.92
4.32
1.89
8.38
27.22
Helichrysum
stuhlmannii
0.92
4.23
1.94
8.2
35.42
Alchemilla
argyrophylla
0.83
3.45
1.32
6.68
42.10
Galium
ruwenzoriense
Group: Kenya & Elgon; average dissimilarity 74.75%; total of 83 species
Dissimilarity/
Contribution %
Species
Avera Averag Average
dissimilarity Std dev.
e
ge
abund abunda
nce 2
ance
1
1.00
0.00
2.01
6.22
2.69
Dendrosenecio
keniodendron
0.92
0.18
1.54
1.71
2.05
Carduus
schimperi
0.73
1.43
1.43
1.91
Koeleria capensis 0.08
0.25
0.91
1.40
1.47
1.88
Helichrysum
formosissimum
0.08
0.64
1.29
1.22
1.73
Geranium
arabicum
0.64
1.25
1.27
1.67
D. elgonensis ssp. 0.00
barbatipes
0.67
0.18
1.23
1.20
1.65
Galium
acrophyum
0.67
0.09
1.23
1.28
1.64
Alchemilla
microbetula
0.33
0.91
1.22
1.27
1.63
Galium
ruwenzoriense
0.36
1.21
1.17
1.61
Sagina afroalpina 0.83
Group: Mt Kenya & Kilimanjaro; average dissimilarity 75.97%; a total of 71 species
1.00
0.00
2.21
4.95
2.91
Dendrosenecio
keniodendron
0.92
0.00
2.03
2.69
2.67
Carduus
schimperi
0.75
0.00
1.64
1.60
2.16
Lobelia telekii
0.00
0.75
1.60
1.63
2.11
Euryops
dacrydioides
0.25
0.92
1.58
1.46
2.08
Erica arborea
0.17
0.83
1.58
1.55
2.08
Helichrysum
newii
0.67
1.40
1.25
1.84
Koeleria capensis 0.08
0.33
1.40
1.21
1.84
Sagina afroalpina 0.83
0.67
0.17
1.38
1.19
1.82
Galium
acrophyum
0.58
0.08
1.31
1.10
1.73
Arabis alpina
Group: Mt Elgon & Kilimanjaro; average dissimilarity 73.25 %; a total of 86 species
Cumulativ
e%
2.69
4.740.08
6.65
8.53
10.26
11.93
13.58
15.22
16.86
18.47
2.91
5.59
7.74
9.85
11.93
14.01
15.85
17.69
19.51
21.23
77
0.08
1.66
2.09
2.27
2.27
Galium ruwenzoriense 0.91
0.75
1.45
1.65
1.98
4.25
Euryops dacrydioides 0.00
0.18
0.83
1.38
1.50
1.88
6.13
Helichrysum newii
0.91
0.25
1.36
1.48
1.86
8.00
Helichrysum
formosissimum
Erica arborea
0.27
0.92
1.36
1.42
1.86
9.85
0.64
0.00
1.24
1.26
1.70
11.55
Dendrosenecio
elgonensis
ssp.
barbatipes
0.64
0.00
1.24
1.27
1.69
13.24
Lobelia telekii
0.00
0.58
1.18
1.13
1.61
14.85
Lobelia deckenii
0.55
0.00
1.11
1.06
1.52
16.37
Artemisia afra
0.55
0.08
1.11
1.04
1.51
17.88
Agrostis gracilifolia
Group: Mt Kenya & Bale; average dissimilarity is 82.59%; a total of 101 species
1.00
0.00
2.09
5.14
2.54
2.54
Dendrosenecio
keniodendron
0.75
0.00
1.55
1.60
1.88
4.41
Lobelia telekii
0.00
0.75
1.54
1.60
1.87
6.28
Dicrocephala
chrysanthemifolia
0.92
0.19
1.54
1.66
1.86
8.14
Carduus schimperi
0.75
0.00
1.51
1.61
1.83
9.97
Anthoxanthum nivale
0.75
0.00
1.51
1.61
1.83
11.81
Alchemilla
argyrophylla
0.00
0.63
1.33
1.22
1.62
13.42
Carex simensis
0.00
0.63
1.25
1.23
1.52
14.94
Festuca simensis
0.00
0.56
1.23
1.09
1.49
16.42
Swertia abyssinica
0.25
0.69
1.22
1.16
1.47
17.90
Colpodium hedbergii
Group: Mt Elgon & Bale; average dissimilarity is 85.27%; total no of species 113 species
0.00
1.71
2.75
2.01
2.01
Galium ruwenzoriense 0.91
0.91
0.13
1.53
1.91
1.80
3.80
Helichrysum
formosissimum
0.09
0.75
1.33
1.46
1.56
5.36
Dicrocephala
chrysanthemifolia
0.64
0.00
1.26
1.27
1.47
6.84
Anthoxanthum nivale
0.00
0.69
1.25
1.42
1.46
8.30
Colpodium hedbergii
0.64
0.00
1.18
1.27
1.39
9.69
Dendrosenecio
elgonensis
ssp.
barbatipes
0.64
0.00
1.18
1.27
1.38
11.07
Lobelia telekii
0.64
0.19
1.14
1.14
1.34
12.41
Geranium arabicum
0.00
0.63
1.14
1.24
1.34
13.75
Festuca simensis
0.18
0.63
1.13
1.14
1.13
15.08
Galium acrophyum
Group: Mt Kilimanjaro & Bale; average dissimilarity 85.19%; total no of species: 104 species
0.83
0.00
1.66
2.04
1.95
1.95
Helichrysum newii
0.92
0.13
1.65
1.88
1.94
3.89
Erica arborea
0.00
1.51
1.62
1.77
5.66
Euryops dacrydioides 0.75
0.08
0.75
1.47
1.45
1.73
7.39
Dicrocephala
chrysanthemifolia
0.67
0.00
1.43
1.33
1.68
9.07
Pentaschistis
78
borussica
0.00
0.69
1.36
1.39
1.60
Colpodium hedbergii
0.00
0.63
1.33
1.22
1.56
Carex simensis
0.67
0.06
1.30
1.27
1.52
Conyza subscaposa
0.17
0.69
1.29
1.22
1.52
Haplocarpha
rueppellii
0.67
0.00
1.29
1.35
1.51
Carduus keniensis
Group: Mt Kenya & Simen; average dissimilarity: 83.10%; total species: 137 species
1.00
0.00
1.54
6.96
1.85
Dendrosenecio
keniodendron
0.00
0.75
1.20
1.67
1.44
Lobelia
rhynchopetalum
0.25
1.00
1.17
1.67
1.41
Colpodium hedbergii
0.75
0.00
1.14
1.66
1.37
Lobelia telekii
0.75
0.00
1.12
1.66
1.35
Anthoxanthum nivale
0.75
0.00
1.12
1.66
1.35
Alchemilla
argyrophylla
0.00
0.75
1.12
1.66
1.35
Festuca simensis
0.00
0.67
1.03
1.38
1.24
Trifolium multinerve
0.00
0.67
1.02
1.37
1.23
T. cryptopodium
0.00
0.67
1.01
1.37
1.22
Rytidosperma
subulata
Group: Mt Elgon & Simen; average dissimilarity: 88.24%; total species: 147 species
0.00
1.00
1.43
7.92
1.63
Colpodium hedbergii
0.00
1.29
1.91
1.47
Galium ruwenzoriense 0.91
Helichrysum
0.91
0.00
1.29
1.92
1.46
formosissimum
0.00
0.75
1.11
1.68
1.26
Lobelia
rhynchopetalum
0.00
0.75
1.06
1.68
1.20
Crassula granvikii
0.00
0.75
1.05
1.67
1.18
Festuca simensis
0.09
0.75
1.00
1.50
1.13
Alchemilla
microbetula
0.00
0.67
0.96
1.39
1.09
Trifolium multinerve
0.00
0.67
0.95
1.38
1.07
Rytidosperma
subulata
0.64
0.00
0.94
1.29
1.06
Anthoxanthum nivale
Mt. Kilimanjaro & Simen; average similarity: 87.98%; total no of 140 species
0.00
1.00
1.53
6.58
1.74
Colpodium hedbergii
0.83
0.00
1.24
2.12
1.40
Helichrysum newii
0.00
0.75
1.19
1.66
1.36
Lobelia
rhynchopetalum
0.00
1.12
1.67
1.27
Euryops dacrydioides 0.75
0.00
0.75
1.11
1.65
1.27
Festuca simensis
0.92
0.25
1.10
1.50
1.25
Erica arborea
0.67
0.00
1.05
1.36
1.19
Pestaschistis
borussica
0.00
0.67
1.02
1.37
1.16
Trifolium multinerve
0.00
0.67
1.02
1.36
1.16
Trifolium
10.67
12.23
13.75
15.27
16.79
1.85
3.29
4.70
6.08
7.43
8.78
10.13
11.37
12.59
13.82
1.63
3.09
4.55
5.82
7.02
8.20
9.33
10.42
11.49
12.56
1.74
3.15
4.50
5.78
7.04
8.30
9.48
10.65
11.80
79
cryptopodium
0.17
0.75
1.01
1.37
1.15
12.96
Alchemilla
microbetula
Group: Mt Bale & Simen; average dissimilarity: 74.34%; total no. of species: 130 species
0.00
0.67
0.99
1.38
1.33
1.33
Trifolium multinerve
Crassula granvikii
0.19
0.75
0.96
1.34
1.29
2.62
0.13
0.67
0.92
1.26
1.24
3.86
Rytidosperma
subulata
0.13
0.67
0.92
1.25
1.23
5.09
Trifolium
cryptopodium
0.19
0.67
0.89
1.20
1.19
6.28
Crepis dianthoseris
0.75
0.33
0.88
1.15
1.18
7.47
Dicrocephala
chrysanthemifolia
0.63
0.17
0.86
1.15
1.18
7.47
Carex simensis
0.25
0.58
0.83
1.06
1.12
9.74
Ranunculus
oreophytus
0.19
0.58
0.81
1.08
1.09
10.83
Carduus schimperi
0.69
0.42
0.77
1.04
0.04
11.87
Haplocarpha
rueppellii
Group: Mt Kenya & Ruwenzori; average dissimilarity 81.67%; total no of species: 68
Dendrosenecio
1.00
0.00
2.44
4.49
2.98
2.98
keniodendron
0.92
0.00
2.44
2.60
2.74
5.72
Carduus schimperi
0.00
0.92
2.21
2.61
2.70
8.42
Dendrosenecio
advinalis
ssp.
advinalis
0.00
0.92
2.20
2.60
2.69
11.12
Helichrysum
stuhlmannii
0.83
0.00
1.97
1.97
2.41
13.52
Sagina afroalpina
0.00
0.75
1.96
1.62
2.40
15.92
Lobelia wollastonii
Alchemilla johnstonii
0.08
0.83
1.82
1.69
2.13
18.15
0.75
0.00
1.80
1.57
2.20
20.36
Lobelia telekii
0.33
1.00
1.66
1.34
2.04
22.39
Cardamine obliqua
0.67
0.00
1.66
1.33
2.03
24.42
Galium acrophyum
Group: Mt Elgon & Ruwenzori; average dissimilarity 81.12 %; total no. of species 87
0.00
0.92
1.97
2.74
2.43
2.43
Dendrosenecio
advinalis
ssp.
advinalis
0.00
0.92
1.97
2.75
2.42
4.86
Helichrysum
stuhlmannii
0.00
0.75
1.74
1.65
2.14
7.00
Lobelia wollastonii
0.73
0.00
1.61
1.53
1.98
8.98
Koeleria capensis
0.27
1.00
1.60
1.53
1.98
10.96
Cardamine obliqua
0.91
0.33
1.43
1.27
1.76
12.72
Helichrysum
formosissimum
0.64
0.08
1.41
1.20
1.74
14.46
Geranium arabicum
0.82
0.33
1.36
1.20
1.68
16.14
Luzula abyssinica
0.64
0.17
1.36
1.15
1.68
17.82
Anthoxanthum nivale
0.08
1.35
1.20
1.67
19.49
Helichrysum forskahlii 0.64
80
Group: Mt. Kilimanjaro & Ruwenzori; average dissimilarity 83.65 %; total no. of species 73
0.00
0.92
2.20
2.53
2.63
2.63
Dendrosenecio
advinalis
ssp.
Advinalis
0.00
0.92
2.19
2.54
2.62
5.24
Helichrysum
stuhlmannii
0.00
0.75
1.95
1.60
2.33
7.57
Lobelia wollastonii
0.92
0.17
1.93
1.73
2.31
9.88
Erica arborea
0.83
0.00
1.91
1.99
2.28
12.17
Helichrysum newii
0.83
1.84
1.65
2.20
14.36
Galium ruwenzoriense 0.08
0.08
1.83
1.74
2.19
16.56
Helichrysum forskahlii 0.83
0.25
1.00
1.83
1.54
2.19
18.74
Cardamine obliqua
0.00
1.74
1.60
2.08
20.83
Euryops dacrydioides 0.75
0.67
0.00
1.68
1.30
2.00
22.83
Pentaschistis
borussica
Group: Mt Bale & Ruwenzori; average dissimilarity 90.06%; total no. of species 100
0.00
0.92
2.07
2.58
2.29
2.29
Dendrosenecio
advinalis
ssp.
advinalis
0.00
0.92
2.06
2.59
2.29
4.58
Helichrysum
stuhlmannii
0.00
0.92
2.04
2.64
2.27
6.85
Alchemilla
argyrophylla
ssp.
argyrophylloides
0.83
1.85
1.94
2.06
8.91
Galium ruwenzoriense 0.00
Alchemilla johnstonii
0.00
0.83
1.85
1.96
2.05
10.96
0.00
0.75
1.83
1.61
2.03
12.99
Lobelia wollastonii
0.75
0.00
1.68
1.56
1.86
14.85
Dichrocephala
chrysanthemifolia
0.31
1.00
1.57
1.36
1.75
16.59
Cardamine obliqua
Haplocarpha
0.69
0.00
1.54
1.37
1.71
18.31
rueppellii
0.63
0.00
1.49
1.22
1.66
19.97
Galium acrophyum
Group: Mt Simen & Ruwenzori; average dissimilarity 89.86%; total no. of species 132
0.00
0.92
1.48
2.86
1.65
1.65
Dendrosenecio
advinalis
ssp.
advinalis
0.00
0.92
1.48
2.87
1.65
3.30
Helichrysum
stuhlmannii
0.00
0.92
1.47
2.89
1.64
4.94
Alchemilla
argyrophylla
ssp.
argyrophylloides
0.83
0.00
1.35
2.06
1.51
6.44
Sagina afroalpina
0.83
1.34
2.06
1.49
7.93
Galium ruwenzoriense 0.00
0.00
0.83
1.33
2.07
1.48
9.42
Alchemilla johnstonii
0.00
0.75
1.29
1.67
1.43
10.85
Lobelia wollastonii
81
2.3.6 Community Description
The cover data of the sub-plots (5 sub–plots × 75 plots) were assessed in terms of percentage
and was used to describe the various Afroalpine vegetation communities prevalent in the
sampled plots from the five vegetation types across all the mountains sampled. The
description of the vegetation communities outlined in table 12 (next page) was based on one
or few dominant species in terms of cover to characterize the vegetation communities. For
each of the vegetation types investigated one or few genera and species dominate the
vegetation community and thus the names of the dominant genera and/or species are used to
describe the vegetation communities. For example in bog vegetation type species of Carex
such as C. monostachya and C. runssoroensis dominate the cover of the bog communities and
thus the vegetation community is described as Carex bog community. However other bog
species such as Haplocarpha rueppellii, Subularia monticola, and Hydrocotyle sibthorpioides
among others are well represented in the bog community. In grassland community, usually
the species of three genera viz: Festuca, Agrostis, and Andropogon are the dominant taxa.
Other grass species are also well represented in grassland communities. These includes:
Koeleria capensis, Pentaschistis borussica, P. pictigluma, Pennisetum humile. Non-grass
species frequently occurring in grassland communities include Alchemilla pedata,
Helichrysum citrispinum and Euryops prostrata among others.
In Dendrosenecio and Alchemilla vegetation communities the dominant species are those
belonging to the two genera Dendrosenecio and Alchemilla respectively. There are no
specific genera or species completely dominating in rock outcrop vegetation type. However
members of the genera such as Helichrysum, Festuca, Pentaschistis, Rytidosperma and
Deschampsia among others co–dominate in one or more mountains (Table 12).
82
Table 12: Summary of Community description from the five Afroalpine vegetation types
Mountain
Habitat/plot No.
Community Description
Elgon
Rock outcrop 1
Festuca abyssinica−Helichrysum dominated rock outcrop community
Elgon
Rock outcrop 2
Helichrysum citrispinum−Agrostis sclerophylla dominated rock outcrop community
Elgon
Rock outcrop 3
Carex simensis−Crassula cf schimperi dominated rock outcrop
Elgon
Rock outcrop 4
Helichrysum citrispinum−Deschampsia flexuosa dominated rock outcrop community
Elgon
Grassland 1
Festuca abyssinica−Agrostis −Koeleria capensis−Pennisetum dominated Grassland
Elgon
Grassland 2
Festuca abyssinica−Geranium arabicum−Koeleria capensis−Agrostis dominated Grassland
Elgon
Grassland 3
Festuca abyssinica−Agrostis−Pennisetum humile dominated Grassland community
Elgon
Grassland 4
F. abyssinica−Pennisetum humile−Colpodium hedbergii−Koeleria capensis dominated G/land
Elgon
Dendrosenecio 1
Dendrosenecio elgonensis ssp. elgonensis−Alchemilla argyrophylla dominated woodland
Elgon
Dendrosenecio 2
D. elgonensis ssp. barbatipes−Alchemilla johnstonii−A. argyrophylla dominated woodland
Elgon
Dendrosenecio 3
D. elgonensis ssp. barbatipes−Alchemilla argyrophylla−A. johnstonii dominated woodland
Ruwenzori
Dendrosenecio 1
Dendrosenecio advinalis ssp. advinalis−Arabis alpina dominated forest community
Ruwenzori
Dendrosenecio 2
D. advinalis var. petiolatus−Luzula johnstonii−Lobelia wollastonii dominated forest community
Ruwenzori
Dendrosenecio 3
Dendrosenecio advinalis dominated forest community
Ruwenzori
Dendrosenecio 4
Dendrosenecio advinalis var. petiolatus−Arabis alpina dominated alpine forest community
Ruwenzori
Ruwenzori
Ruwenzori
Ruwenzori
Bog 1
Bog 2
Bog 4
Alchemilla 1
Carex runssoroensis−Hydrocotyle sibthorpioides−Helichrysum formosissimum bog community
Carex runsoriensis−Helichrysum stuhlmannii−Isolepis fluitans dominated bog community
Carex runssoroensis−Alchemilla johnstonii dominated bog community
Alchemilla argyrophylla ssp. argyrophylloides−A. johnstonii dominated shrubland community
83
Ruwenzori
Ruwenzori
Ruwenzori
Kilimanjaro
Kilimanjaro
Kilimanjaro
Kilimanjaro
Kilimanjaro
Kilimanjaro
Kilimanjaro
Kilimanjaro
Kilimanjaro
Kilimanjaro
Kilimanjaro
Kilimanjaro
Bale
Bale
Bale
Bale
Bale
Bale
Bale
Bale
Bale
Bale
Bale
Bale
Bale
Bale
Bale
Bale
Alchemilla 2
Alchemilla 3
Alchemilla 4
Grassland 1
Grassland 2
Grassland 3
Grassland 4
Bog 1
bog 2
bog 3
bog 4
Rock outcrop 1
Rock outcrop 2
Rock outcrop 3
Rock outcrop 4
Rock outcrop 1
Rock outcrop 2
Rock outcrop 3
Rock outcrop 4
bog 1
bog 2
bog 3
bog 4
Grassland 1
Grassland 2
Grassland 3
Grassland 4
Alchemilla 1
Alchemilla 2
Alchemilla 3
Alchemilla 4
Alchemilla johnstonii−A. argyrophylla dominated sclerophyllous shrubland community
Alchemilla argyrophylla ssp. argyrophylloides−A. johnstonii−A. triphylla dominated shrubland community
Alchemilla argyrophylla ssp. argyrophylloides dominated Alchemilla shrubland community
Poa leptoclados−A. microbetula−Festuca abyysinica dominated Grassland community
Festuca abyssinica−Helichrysum citrispinum dominated Grassland community
Festuca abyssinica −Deschampsia flexuosa dominated Grassland community
Pentaschistis/Andropogon TZ 0837 dominated Grassland community
Carex monostachya dominated bog community
Carex monostachya dominated bog community
Carex monostachya−Alchemilla johnstonii dominated community
Carex monostachya−Alchemilla johnstonii dominated bog community
Deschampsia flexuosa−Helichrysum citrispinum−H. newii dominated rock outcrop community
Pentaschistis borussica−Helichrysum forskahlii dominated rock outcrop community
Helichrysum newii−Alchemilla johnstonii dominated rock outcrop community
Helichrysum forskahlii−Pentaschistis borussica−Isolepis dominated rock outcrop community
Festuca cf macrophylla−Helichrysum citrispinum−Pentaschistis pictigluma dominated rock outcrop
Helichrysum citrispinum var citrispinum−Cineraria deltoidea−Crassula dominated rock outcrop
Festuca simensis−Pentaschistis pictigluma−Cineraria deltoidea−Crassula dominated rock outcrop
Carex simensis−Helichrysum citrispinum−Lobelia rhynchopetalum dominated rock outcrop
Carex monostachya−Galium acrophyum (ET−0014)−Deschampsia caespitosa dominated bog community
Carex monostachya−Deschampsia caespitosa−Galium acrophyum dominated bog
Carex monostachya−Haplocarpha rueppellii−Eriocaulon ET−0517 dominated bog
Carex monostachya−Subularia monticola−Alchemilla pedata dominated bog
Festuca simensis−Alchemilla pedata dominated grassland community
Festuca simensis−Euryops prostratus−Alchemilla pedata dominated grassland
Andropogon cf lima−Festuca abyssinica−Alchemilla pedata dominated grassland
Andropogon cf lima−Festuca simensis−Alchemilla pedata−Trifolium elgonensis dominated grassland
Alchemilla haumannii (84%)−Alchemilla cf microbetula dominated Alchemilla community
Alchemilla haumannii (91%) dominated Alchemilla community
Alchemilla pedata (94%) dominated Alchemilla community
Alchemilla microbetula−Alchemilla pedata−Isolepis fluitans dominated Alchemilla community
84
Simen
Simen
Simen
Simen
Grassland 1
Grassland 2
Grassland 3
Grassland 4
Festuca simensis−Pentaschistis pictigluma−Andropogon dominated Grassland community
Festuca simensis (64%)−Alchemilla abyssinica−Pentaschistis pictigluma dominated community
Helictotrichon elegantum−Festuca simensis dominated grassland
Festuca simensis−Carex monostachya−H. citrispinum−Pentaschistis pictigluma dominated grassland
Simen
Rock outcrop 1
Simen
Simen
Rock outcrop 2
Rock outcrop 3
Simen
Simen
Simen
Simen
Simen
Kenya
Rock outcrop 4
bog 1
bog 2
bog 3
bog 4
Rock outcrop 1
Mt. Kenya
Mt. Kenya
Rock outcrop 2
Rock outcrop 3
Rock outcrop 4
Mt. Kenya
Mt. Kenya
Mt. Kenya
Mt. Kenya
Dendrosenecio 1
Dendrosenecio 2
Dendrosenecio 3
Dendrosenecio 4
Mt. Kenya
Bog 1
Mt. Kenya
Mt. Kenya
Mt. Kenya
Bog 2
Bog 3
Bog 4
Colpodium hedbergii−Plantago afra−Pentaschistis pictigluma dominated rock outcrop community
Plantago afra−Cotula abyssinica−Alchemilla microbetula−Gnaphalium unionis−Vulpia bromoides dominated
rock outcrop
Colpodium hedbergii (7%)−Rytidosperma subulata dominated rock outcrop
Rytidosperma subulata−Vulpia bromoides−Crepis rueppellii−Lychnis abyssinica−Pentaschistis pictigluma
dominated rock outcrop
Carex monostachya (60%)−Alchemilla abyssinica−Pentaschistis pictigluma dominated bog community
Cotula cryptocephala−Isolepis fluitans−Ranunculus oligocarpus dominated bog
Carex monostachya−Haplocarpha rueppellii−Ranunculus oligocarpus−Cotula abyssinica dominated bog
Cotula cryptocephala−Callitriche oreophila−Veronica anagalis−aquatica dominated bog community
Dendrosenecio keniodendron−Festuca abyssinica−Pentaschistis borussica dominated rock outcrop community
Festuca abyssinica−Dendrosenecio keniodendron−Alchemilla microbetula−Crepis dianthoseris dominated rock
outcrop
Lobelia telekii−Agrostis−Dendrosenecio keniodendron dominated rock outcrop
No data
Dendrosenecio keniodendron−Festuca pilgeri−Senecio cf polyadenus dominated Dendrosenecio woodland
community
Dendrosenecio keniodendron−Festuca pilgeri−Lobelia telekii dominated Dendrosenecio woodland
No data
Dendrosenecio keniodendron−Alchemilla johnstonii−Festuca pilgeri dominated Dendrosenecio community
Carex monostachya (32%)−Festuca pilgeri−Haplocarpha rueppellii−Ranunculus oligocarpus dominated bog
community
Alchemilla microbetula−Lobelia gregoriana subsp−gregoriana−Festuca pilgeri−Haplocarpha rueppellii
dominated bog
Carex monostachya (61%)−Ranunculus oreophyton dominated bog
Festuca pilgeri−Alchemilla microbetula−Ranunculus stagnalis−Carex monostachya dominated bog
85
2.4 DISCUSSION AND CONCLUSION
2.4.1 Species dominance in the Afroalpine vegetation of Eastern Africa
Compared to other ecosystems such as tropical rainforests and the adjacent Afromontane
moist forest of African Mountains, the Afroalpine zone are considered to be species ‘poor’
(Hedberg, 1970; White, 1978a). Afroalpine vegetation communities of Eastern Africa are
dominated by six families, namely Asteraceae (69 species), Poaceae (37 species),
Caryophyllaceae (16 species), Lamiaceae (12 species), Cyperaceae (11 species) and Rosaceae
(10 species) that constitute 56% of all the species sampled. This indicates the dominance of
few families in the colonization and occurrence of their species in the sampled vegetation
communities and mountains. Asteraceae and Poaceae alone account for more than a third
(38%) of all the species of Afroalpine revealing the importance and dominance of the species
of these two families.
A number of factors probably contributed to the success of these two families. Members of
the grass family for example have tiny light seeds (morphological adaptation) capable of
being blown for a long distance from the source region and presumably deposited in the
virgin non−inhabited bare rocks of the mountains during and after the initial processes of
orogeny (mountain formation). Wind dispersal is one of the main mechanisms effecting long
distance dispersal for Afroalpine plants (Troll, 1952; Coe, 1967; Hedberg, 1964b & 1970).
This is especially true for the light seeded plants like grass species, members of Orchidaceae
and Pteridophytes among others. Moreover, anemochory is thought be the hallmark of
pioneer vegetation (Van der Pijl, 1969) and its significance is directly proportional to the
altitudinal height of the mountains (Hedberg, 1970). A classical circumstantial evidence of
the success of Poaceae and/or members of Asteraceae was revealed by the recent discovery of
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species identified to be Senecio purtschelleri Engl., and Crepis Dianthoseris both members of
Asteraceae at the summit of Mt Kilimanjaro which might be partly explained by recent case
of wind dispersal (Hedberg, 1970).
In addition to having relatively light seeded plants many species of the genus Helichrysum,
Senecio and other genera of Asteraceae that are evidently abundant in Afroalpine plant
communities are morphologically endowed with parachute like structures. These are adapted
for air floatation/dispersal and are capable of being blown high up especially during strong
winds prevalent in Afroalpine zone and storms like cyclones which is not infrequent in high
mountains (Hedberg, 1970). Increased chances of successful establishment of the diaspores
after arrival in the various mountain enclaves (refugia) could have allowed the species and
genera of these dominant families to establish successful populations. This is often possible
through physiological and/or genetic adaptation such as increased power of germination as
well as ability to withstand the vagaries of extreme climatic conditions and biotic competition
for dearth resources especially during the initial colonization of virgin habitats.
The occurrence of fire which may be naturally induced e.g. by volcanicity, lightening, etc. or
mostly caused by anthropogenic factors is rampant in African high Mountains and else
where. Fire is thought to significantly affect the sub−alpine (ericaceous) and alpine plant
communities mostly aiding the spread and dominance of grassland communities (Wesche et
al., 2000). Wesche noted the remarkable ability of grassland communities for their adaptation
to these fires through quick recovery as well as the non−effect of the fire on species
composition and the relative cover of the grass and other associated species while recovering.
This enables grass species and associated herbaceous species to expand their ranges through
rapid regeneration after incidents of fire. This explains the frequent occurrence and relatively
87
high cover of grass and associated species not only in the grassland vegetation but also in
other Afroalpine vegetation types especially in rock outcrop. Hence grassland is the most
conspicuous vegetation type in terms of percentage land cover in the entire mountain studied.
Mount Rwenzori is an exception to this pattern since it is dominated by extensive bogs (in the
valleys e.g. Bujuku) and Dendrosenecio forest (slopes and ridges) due to the exceptionally
high precipitation.
2.4.2 Description of the communities of the five vegetation types of Afroalpine zone
The communities frequently occurring in the five vegetation types sampled, viz., rock
outcrops, bogs, grasslands, Dendrosenecio forest/woodland and Alchemilla shrubland
correspond well with those described by Hedberg (1964b). The main communities prevalent
in these vegetation types include Carex runssoroensis−C. monostachya−Deschampsia
caespitosa dominated bogs. The genera Carex and Deschampsia are the typical bog species
that usually requires standing water or soil saturated with water common in this vegetation
type. In most mountains studied, C. monostachya is the common dominant species in bogs
except in Mt Rwenzori which, is dominated by C. runssoroensis. Other bog species common
in this vegetation type are Haplocarpha rueppellii, Hydrocotylee sibthorpioides and
Subularia monticola among others.
In grassland communities, Festuca species often form the dominant cover in most of the
grasslands studied. Festuca abyssinica and F. simensis are the two main species with the
former being the more widespread species. Other genera that are very common in grassland
vegetation type include Pentaschistis (represented mainly by P. borussica and P. pictigluma),
Andropogon (A. lima), Agrostis (A. gracilifolia and A. schimperiana), Pennisetum (P. humile)
and Poa (P. leptoclados). In Dendrosenecio forest/woodland vegetation community the
88
dominant species are the woody Dendrosenecio species in the upper story and Alchemilla
species in ground cover. Usually in each mountain an endemic or near endemic species of
Dendrosenecio occur. For example in Mt Elgon it is represented by D. elgonensis while in Mt
Rwenzori and Mt Kenya it is represented by D. advinalis and D. keniodendron species
respectively. In Alchemilla vegetation type it is usually dominated by the genus Alchemilla
represented mostly by Alchemilla argyrophylla, A. johnstonii, A. pedata and A. haumannii
species. Although species of Alchemilla are frequent across the entire mountains, Alchemilla
vegetation community type is not well developed as only two plots in Bale and Rwenzori
could be found for analysis. In rock outcrop vegetation type several genera including
Helichrysum, Festuca, Koeleria, Colpodium, Plantago and Pentaschistis co–dominate the
rock outcrop communities. The species that forms the dominant cover of the above genera
include: Helichrysum citrispinum, H. forskahlii, H. newii, Festuca simensis, F. abyssinica,
Koeleria capensis, Plantago afra, Pentaschistis pictigluma and P. borussica. Other species
that are frequent in rock outcrop community are Cineraria deltoidea, Deschampsia flexuosa
and Crassula species.
2.4.3 Comparative assessment of species composition between and within five vegetation
types of Afroalpine zone of Eastern Africa
Among the 20 plots of rock outcrop sampled across the six mountains reveal marked
similarities between all the mountains except Simen in terms of the structure of communities
and species assemblage/composition. The dominant genera and species in this vegetation type
that are at least prevalent in two or more mountains are Helichrysum represented by H.
citrispinum in Mt Elgon, Kilimanjaro, and Bale. In addition H. newii and H. forskahlii occur
in Mt Kilimanjaro. The above mountains occur along the eastern side of the rift valley.
Festuca is represented by F. abyssinica in Mt Kenya and Elgon, which are geographically in
89
close proximity thus explaining the observed pattern of similarity while F. simensis and F.
macrophylla occur in Bale. Three species of the genus Deschampsia occur in Afroalpine zone
of Eastern Africa. The first one D. caespitosa is a bog species while D. flexuosa var.
afromontana and D. angusta are common in rock outcrop community in most of the
mountains but forms dominant cover in Mt Elgon and Kilimanjaro rock outcrop community.
The genus Pentaschistis is well represented in rock outcrop community by two species viz. P.
borussica in that it forms dominant cover in Mt Kilimanjaro, and P. pictigluma var.
pictigluma with golden colour inflorescence common in Bale and Simen. This variety is
endemic to Ethiopian Mountains. The Alchemilla species prevalent in rock outcrop
community include A. microbetula in Mt. Kenya and Simen, A. johnstonii in Kilimanjaro and
A. argyrophylla in Mt. Kenya and Ruwenzori. However in Mt Kenya and Rwenzori are
represented by two different subspecies of A. argyrophylla (A. argyrophylla ssp.
argyrophylla in Mt Kenya and A. argyrophylla ssp. argyrophylloides in Rwenzori).
Carex species are predominantly bog species but occasionally is found in other vegetation
types such as rock outcrop community especially in wet zones. For example C. simensis
occur in Elgon and Bale both of them being relatively old mountains along the eastern rift
valley. Agrostis species are among the dominant genera in grassland community but it
frequently occurs in rock outcrop community e.g. Agrostis sclerophylla occur in the mountain
summit of Mt. Elgon & Kenya. Crassula species are often found in crevices of rock outcrop
community e.g. C. schimperi occur both in Elgon and Bale Mountain. Lobelia is among the
few woody species prevalent in Afroalpine zones of Eastern Africa. In rock outcrop
community, L. telekii frequently occur in Mt Kenya while L. rhynchopetalum is common in
Bale. Crepis is represented by C. dianthoseris in Mt Kenya and C. rueppellii in Simen.
Cineraria deltoidea is another common rock outcrop species that occur in several mountains
90
especially common in Simen and Kilimanjaro. Other dominant genera in the rock outcrop
community which, are mountain specific include Rytidosperma (R. subulata), Colpodium (C.
hedbergii), Plantago (P. afra), Cotula (C. abyssinica), Gnaphalium unionis, Vulpia
bromoides, Crepis rueppellii and Lychnis abyssinica all in Simen; Dendrosenecio
keniodendron in Mt Kenya and Isolepis sp. in Kilimanjaro.
In boggy vegetation type there was generally uniformity across all the mountains in the
generic composition. However there were differences in the species composition, which,
were dominated by hydrophilic species mainly from the tussock forming genus Carex and
acaulecent plants like Haplocarpha species. Mount Ruwenzori, the wettest mountain
(Hedberg; 1964b; Wesche et al., 2000) significantly differs from the rest of the mountains in
terms of species composition of the most dominant taxa. For example the tussock forming
Carex runsoroensis is the dominant species in all the four vegetation types sampled from
Rwenzori with almost pure stands of this species in some cases in terms of cover. The rest of
the mountains are dominated by Carex monostachya in association with various members of
other genera. In Ruwenzori, beside Carex, other taxa with significant representation include
Hydrocotyle sibthorpioides, Isolepis fluitans, Helichrysum stuhlmannii, H. formosissimum
and Alchemilla argyrophylla ssp. argyrophylloides (Rwenzori endemic). In Mt Kilimanjaro,
C. monostachya and Alchemilla johnstonii co−dominates in the bog community.
In the remaining three mountains viz. Bale, Mt Kenya and Simen other dominant genera
include Ranunculus variously represented by R. oligocarpus in Mt Kenya and Simen, R.
oreophytus and R. stagnalis in Mt Kenya. Haplocarpha is represented by H. rueppellii in Mt
Kenya, Bale and Simen while Alchemilla is represented by A. microbetula in Mt Kenya, A.
pedata in Bale and A. abyssinica in Simen. However, Cotula cryptocephala, Isolepis fluitans,
91
Callitriche oreophila, Veronica anagalis−aquatica are other dominant species in Mt Simen
only although some of them may have been recorded from the other mountains. This makes
Mt Simen rather unique compared to other mountains. From the generic composition, it is
clear that the water loving plants (hydrophyllics taxa) or bog species dominate since water is
critical in determining distribution and dominance of species in this vegetation type as the
Afroalpine species exhibit different levels of water tolerance. Consequently the wettest
mountain Rwenzori, which receives rain almost every day, has the best developed bogs with
almost pure stands of tall lash green Carex runssoroensis tussocks making it significantly
different from the rest of the mountains. Often the bogs here are characterized by the
permanent presence of surface water in most parts of the expansive bogs.
The genera and species of the family Poaceae dominate the floristic composition of grassland
community. In almost all the mountains, species of Festuca (F. abyssinica in EA Mountains
and F. simensis in Ethiopian Mountains) dominates the species composition of the sampled
plots. In Mt. Elgon, F. abyssinica, Koeleria capensis, Pennisetum humile and Colpodium
species forms the dominant taxa in grassland communities. In Mt. Kilimanjaro, Poa
leptoclados, F. abyssinica, P. borrusica and Andropogon lima forms the dominant species
with Alchemilla microbetula having significant occurrence. Festuca simensis, P. pictigluma,
Andropogon lima and Helictotrichon elegantum dominate grassland species in Simen with
significant presence of A. abyssinica and H. citrispinum species. In Bale, F. simensis, A.
pedata and A. lima forms the dominant species of grassland community.
Species of the genus Pentaschistis are among the most conspicuous grass species in the
Afroalpine zone along with Festuca species. The species of Pentaschistis that are known to
occur in the study area and, which have been sampled both within and outside the plots
92
include P. borussica, P. trisetoides, P. dolichochaeta, P. chrysurus, P. pictigluma var.
pictigluma, P. pictigluma var. minor, P. pictigluma var. gracilis, and P. pictigluma var.
mannii. Pentaschistis borussica is the most widely distributed species occurring in all the
mountains except Simen. Its preferred vegetation communities are rock outcrop and
grassland, rarely occurring in bogs and Dendrosenecio forest/woodland. The other widely
distributed species is P. pictigluma. However this species was represented in most of the
mountains by one or two varieties. Pentaschistis pictigluma var. pictigluma and P. trisetoides
are endemic to Ethiopian Mountains. Most of the known collections including collections
from this study of P. pictigluma var. gracilis are from Ethiopia. In addition P. dolichochaeta,
a robust species is endemic to Shewa region of Ethiopia especially in Wafwasha forest and
the nearby Ancober highland occurring below Afroalpine zone [sub– alpine] (2900 to 3000
m). Similarly P. chrysurus is endemic to Tanzania (Mt. Kilimanjaro, Mt. Meru and nearby
highlands including Mt. Hanang). This species also occurs below the alpine zone (2500−3400
m) mostly in thickets around the forest zone.
In Dendrosenecio vegetation type, only three East Africa countries viz., Mt Kenya, Elgon
(Kenya/Uganda) and Rwenzori were sampled as this conspicuous Afroalpine genus was
absent from Ethiopian Mountains. Mount Kilimanjaro, the driest of all the mountains studied
had only scattered Dendrosenecio trees in the Afroalpine zone, which could not form
continuous plot of Dendrosenecio community comparable to other mountains for sampling
and analysis. Hence no continuous woodland/forest plots of the required size of
Dendrosenecio could be found in Mt. Kilimanjaro.
Each of the three mountains sampled had an endemic vicariant species of Dendrosenecio as
the dominant species. This was associated frequently and/or co−dominated occasionally with
93
other woody (Lobelia) or shrubby Alchemilla or Helichrysum species. For example, in Mt
Elgon D. elgonensis (D. elgonensis ssp. elgonensis and D. elgonensis ssp. barbatipes) was
the dominant species frequently co−dominated at the ground level by A. argyrophylla ssp.
argyrophylla and A. johnstonii. Dendrosenecio keniodendron dominates Mt Kenya
Dendrosenecio woodland with frequent association at the ground level with Festuca pilgeri
and occasionally A. johnstonii. In Rwenzori, Dendrosenecio vegetation type was well
developed often forming dense moist forest dominated by D. advinalis ssp. advinalis and D.
advinalis var. petiolatus with branches and dead Dendrosenecio wood on the ground covered
with thick moss and other epiphytic species like Galium ruwenzoriense. At the ground level
this species is frequently associated with Arabis alpina, Lobelia wollastonii (Rwenzori and
Virunga endemic) and Luzula johnstonii.
Alchemilla community was the least extensively developed vegetation type in the whole of
the study area although well represented by many species across the entire mountains. As
such only eight plots were sampled from two mountains (Bale in Ethiopia and Rwenzori in
East Africa) and the other mountains lacked sizable plots for sampling and analysis. In
Ruwenzori the Alchemilla communities were dominated in terms of species cover and
presence by A. argyrophylla ssp. argyrophylloides and A. johnstonii with frequent association
of A. triphylla. On the other hand, A. haumanii was basically the sole dominant species in
Bale often with cover of over 84%. In one plot however, A. micrbetula, A. pedata and
Isolepis fluitans (all frequently found in bogs) dominated the Alchemilla community (was
perhaps wet zone resembling boggy conditions thus attracting nearby bog species).
Jaccard’s pair wise similarity index revealed that rock outcrop and grassland vegetation types
were the most similar with J’ value of 52. A total of 118 species were common from these
94
two vegetation types while they have combined differences of 109, that is, either occurs in
rock outcrop or grassland. The other pairs that share relatively higher degree of similarity are
bogs and grassland (J’=44.0; common−96; different 122) and rock outcrop and bog (J=41.0;
common−101; different−143). The higher degree of similarity can be attributed to a number
of factors. First the most important abiotic factors affecting the Afroalpine species include
(but not limited to) light, water and edaphic factors respectively in addition to intra and
interspecific competition. One common thing with these three vegetation types was the
openness and the availability of maximum light due to the near absence of large woody
species of Dendrosenecio, Lobelia and shrubby and dense Alchemilla/Helichrysum species
that have the potential of obstructing/and limiting light penetration to the ground layer. Hence
the genera and species common to these vegetation communities are non−shade loving
herbaceous taxa often with creeping, climbing, tussock forming and acaulecent habits. As a
result there are marked dissimilarity between grassland and Dendrosenecio communities
(J=22.0; common−40; different−145) and/or rock outcrop and Alchemilla (J=26.0;
common−55; different−145).
Both Dendrosenecio and Alchemilla communities, which are dominated by the species of
these two genera, have the tendency to fairly give dense cover to the ground below
significantly limiting light penetration on to the ground layer. This limit the establishment or
occurrence of many Afroalpine species–mostly herbaceous annuals with creeping habit,
reduced stems and non−tussock forming grass species. In most cases these constitute
significant number of Afroalpine species (Hedberg, 1964b). Therefore Dendrosenecio and
Alchemilla vegetation types are thought to be the most species poor among all the five
vegetation communities sampled during the study.
95
In addition to its primary role of photosynthetic regulation, light from the solar radiation has
cascading effect on other equally important abiotic factors such as its direct role of
temperature regulation as well as its influence on evapotranspiration, frequent thawing
(daytime) and freezing (night) peculiar in Afroalpine zone which creates “artificial” water
shortage (Hedberg, 1964b; Wesche, 2000). This is essentially why there are large fluctuations
of temperature in the Afroalpine zones of the high mountains in tropical Africa during the
day and night, that is, “summer everyday and winter every night” (Hedberg, 1957). Sun light
too influences directly water uptake by plant since during the night and the early hours of the
day, the soil temperatures is below zero, impairing water and nutrient uptake or at least
slowing the process of water uptake significantly. Consequently most species found in these
three vegetation types, which are species rich have developed both physiological and
morphological adaptation to tide over the extreme conditions of temperature fluctuations and
artificial water shortages. These include developing xeromorphic characteristics like stem
reduction e.g. Haplocarpha species and Haplosciadium species, tussock forming habit for
insulation and protection of vascular tissue e.g. Carex and Festuca species, and
sclerophyllous shrubby characters with convolute or revolute leaves e.g. Alchemilla and
Helichrysum species to reduce evapotranspiration (Hedberg, 1964b).
2.4.4 Comparative assessment of species richness and abundance between the five
vegetation types
In terms of species richness, the rock outcrop vegetation type was the most species rich and
significantly different with regard to this parameter compared to both Dendrosenecio and
Alchemilla vegetation communities. Similarly bogs are more species rich compared to
Dendrosenecio vegetation community. Hence the null hypothesis of no significant
differences in the plant species richness between the vegetation types is rejected (P< 0.05). A
total of 188, 157, 68 and 77 species were either recorded or collected from rock outcrops,
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bogs and grasslands, Dendrosenecio and Alchemilla vegetation types respectively. A number
of possible reasons can be given to explain the richness of the rock outcrop compared to other
vegetation types. First, rock outcrop provides many microhabitats or niches where diverse
number of species with different requirements of optimal conditions of abiotic and biotic
factors can thrive compared to other vegetation types. For example in rock outcrops we have
caves for shade lovers and epiphytic plants, crevices and shallow depressions where water
can collect to mimic boggy condition where bog species can colonize and shallow or thin soil
cover that favours grasses and herbaceous species. However species diversity and evenness
between the five vegetation types appear to be uniform with no significant variations.
2.4.5 Comparative assessment of species composition between the alpine zone of the six
mountains of Eastern Africa
Comparative analysis of the species composition and community ecological analysis revealed
a general pattern of similarity and/or differences between the six mountains based on several
factors: −
•
Geographical location of the mountains
•
Age of the formation of the mountains
•
Geological history of the mountains
•
Climatic factors
•
Edaphic or Soil factors
From this study, the degree of the similarity between the mountains was more or less
inversely related to geographical distance between the mountains and interestingly their
location along the rift valley systems (eastern verse western arm). The “sky−islands” of each
of the mountains tops serves as refugia with effective isolation similar to oceanic islands off
the coast of the nearest mainland. The only biotic interchange between these enclaves is
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thought to be almost always through long distance dispersal followed by successful
establishment of the migrating/colonizing species (Hedberg, 1970). Hence it follows that “the
sky islands” isolations increases with both distances from each other and the height of
individual mountain. Therefore, the distances (geographical location) as well as the location
along the arm of rift valley systems (Rwenzori and Simen on the western arm and East Africa
Mountains and Bale on the eastern arm) influence species composition of the Afroalpine flora
across the mountains.
Consequently Mt Kenya, Kilimanjaro, and Elgon share many common species and are
floristically similar compared to the two Ethiopian Mountains and to a lesser extent
Ruwenzori since the latter has unique attributes including its location on the western arm of
the rift valley system and non–volcanic origin. Mount Kenya and Kilimanjaro, the two
closest mountains in East Africa show the highest degree of similarity (Jaccard’s similarity
index: 44.4). In addition the similarities between the East Africa mountains viz: Mt Kenya,
Kilimanjaro, Elgon and to a lesser extent Mt Ruwenzori is due to having similar species
composition especially such species like Anthoxanthum nivale K. Schum., Alchemilla
argyrophylla Oliv., Carduus keniensis R.E.Fr., Helichrysum newii Oliv. & Hiern among
others that frequently occurred in many plots across these mountains. However each of the
mountains has its endemic species which are localized to that mountain. For example,
Dendrosenecio elgonensis (T.C.E.Fr.) E.B. Knox is endemic to Mt Elgon, Euryops
dacrydioides Oliv. (Mt Kilimanjaro), Dendrosenecio keniodendron (R.E.Fr. & T.C.E.Fr.)
B.Nord. (Mt Kenya), D. keniensis (Baker f.) Mabb. (Kenya), Helichrysum stuhlmannii O.
Hoffm., D. advinalis (Stapf) E.B. Knox and Lobelia wollastonii Baker f. that are localized to
Mt Ruwenzori.
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Similarly Bale and Simen have higher Jaccard’s similarity index than between any other pair
for the same reason (J’ 47.05). Again Bale and Simen (Ethiopian) share many species that are
endemic to these two mountains. These species include Rytidosperma subulata (A.Rich.)
Cope, Lobelia rhynchopetalum Hemsl, Festuca simensis A. Rich., Pentaschistis pictigluma
(Steud.) Pilg. (including var. pictigluma and var. gracilis) and Valerianella microcarpa
Loisel among others. Mount Bale and Simen like the East Africa Mountains have their own
endemic species such as Rytidosperma grandiflora (Hochst. Ex A. Rich.) S.M. Phillips,
Aphanes bachitii (Haum. & Balle) Rothm., Pimpinella pimpinelloides (Hochst.) Wolff and
Argyrolobium schimperianum Hochst. Ex A.Rich. for Simen. Swertia crassiuscula ssp.
robusta Sileshi, S. macrosepala ssp. microseperma Sileshi, Geranium sp. = Miehe 3002
(Bale) and Euryops prostrata B. Nord. are endemic to Bale. Hence there are various levels of
separation between the mountains in terms of the species composition from the 75 plots
sampled.
Mount Ruwenzori exhibit marked difference from other East Africa and Ethiopian Mountains
in that it has a number of species endemic to this mountain and was different from the rest of
the mountains. This was the only mountain in East Africa sampled that is non−volcanic, i.e. it
was formed by the up thrust (rise) of part of the African plate and consequently the soil and
the parent rocks are quite different from those of East Africa. Secondly Rwenzori was the
wettest and presumably the coldest mountain sampled. Although it has dense and moist
Dendrosenecio forest, thick Alchemilla community and extensive swampy bogs it’s the most
species poor compared to the rest of the mountains. The same reasons or factors such as
sunlight availability (due to dense growth and cloudiness), water stress etc discussed earlier
under the vegetation communities can be stated to explain this observed pattern.
99
Ruwenzori was also the only mountain after Simen that is located in the western arm of rift
valley, which therefore provides another third level of isolation (barrier to dispersal) since the
rift systems provides partial isolation between the mountains (Hedberg, 1970). The parent
rock and soil types of Rwenzori was significantly different from all other mountains studied
since it constitute ancient Eozoic rocks (gneises and schists) that is, metamorphic rocks
including such rocks like mica−schist, amphibolites, calc−schists, quartzites, muscovites and
cordierite−schists (Gregory, 1921) while the rest of the mountains were formed from basaltic
lava.
The species richness among five out of the six mountains (Kenya, Elgon, Rwenzori,
Kilimanjaro and Bale) is not significantly different from each other. Consequently the null
hypothesis (hypothesis two) is accepted, that is, p> 0.05. However the species richness of
Simen mountain is significantly different from the rest (p< 0.05) thus in this case rejecting the
null hypothesis.
The age of the six mountains studied ranges from 2 to 3.5 MY BP to over 20 MY BP in the
case of Mt Elgon and Ethiopian Mountains. The Ethiopian massifs started rising about 70
MY BP and are therefore much older than those of East Africa. Consequently they have had a
considerable geological time scale for accumulation of divergent characters and evolution of
diverse Afroalpine flora compared to say Mt Kenya and Kilimanjaro (“baby Mountains”).
One other major difference between Ethiopian Mountains and those of East Africa was the
absence of keystone genus of the Afroalpine flora, the giant rosette plant viz: the genus
Dendrosenecio (Asteraceae) and the vicariant nature of the species of the genus Lobelia
(Lobeliaceae). Moreover, Simen and Bale exhibit different rainfall pattern in that they have
unimodal pattern of rainfall distribution where there is single long rainy season and single
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short dry spell compared to East African mountains which have bimodal pattern of rainfall
(two rainy seasons and two dry seasons). These factors perhaps explain the species richness
of these mountains and the high degree of dissimilarity with those of East Africa (Gashaw &
Fetene, 1996).
Despite the above stated differences between the mountains regarding the geological history,
climatic condition variations and existence of differential characteristics of the edaphic
factors the overall similarities in the species composition between the mountains is quite
remarkable. Hence there are no significant differences in the species composition among the
mountain studied thus affirming the statement of the hypothesis.
2.4.6 Heterogeneity Assessment
The mean values of R for the ANOSIM results indicate that the five vegetation types were
significantly different from each other and can be easily recognized thus rejecting the null
hypothesis four (all the vegetation types of Afroalpine zone are homogenous). However the
heterogeneity of the vegetation types was blurred by overlap in terms of species composition.
For example vegetation types such as rock outcrop vs. grassland are barely separable at all
(R< 0.25) while rock outcrop vs Dendrosenecio, rock outcrop vs Alchemilla, bog vs
Alchemilla, bog vs grassland, grassland vs Alchemilla, and Dendrosenecio vs Alchemilla are
overlapping but fairly separable. However, rock outcrop vs bog and bog vs Dendrosenecio
are overlapping but clearly separable (R > 0.5).
In the SIMPER analysis among the five vegetation types− rock outcrops, bogs, grasslands,
Dendrosenecio and Alchemilla −from the entire six mountains communities several keystone
species that are ubiquitously distributed emerge that greatly contribute to the overlapping of
101
the vegetation communities. Key among these species are: Erica arborea (Ericaceae),
Colpodium hedbergii (Poaceae), Helichrysum formosissimum (Asteraceae), Cerastium
octandrum (Caryophyllaceae), Gallium acrophyum (Rubiaceae), Anthoxanthum nivale
(poaceae), Alchemilla argyrophylla (Rosaceae) and Arabis alpina (Brassicaceae) which
frequently occur in the various vegetation types studied.
The species richness, abundance and diversity varies across the vegetation communities and
mountains. Species composition among the five vegetation communities investigated show
higher similarities between grassland, rock outcrop and bog communities compared to
Dendrosenecio and Alchemilla communities. The former also show higher species diversity
and richness compared to the latter. Mountains from East Africa (Kenya, Tanzania and
Uganda) countries show less species richness and less diversity compared to the Ethiopian
mountains. Species composition among the mountains of East Africa show higher degree of
similarity compared to the Ethiopian mountains. Factors such as the age, geological history of
the mountains, distance between the mountains, edaphic factors and climatic conditions are
responsible for the observed patterns of species composition among the various mountains
investigated.
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CHAPTER THREE: PHENETIC AND PHYLOGENETIC ANALYSIS OF THE
GENUS PENTASCHISTIS (NEES) SPACH (POACEAE) OCCURRING IN EASTERN
AFRICAN MOUNTAINS
3.1 LITERATURE REVIEW
3.1.1 Distribution and Description of the genus Pentaschistis
The African endemic genus Pentaschistis is among the largest genera in the tribe
Danthonieae (Danthonioideae, Poaceae). There are about 70 species of the genus
Pentaschistis, mainly occurring in Southern Africa but also extending northwards on the high
mountains of East Africa, Ethiopia, Cameroon and Madagascar [figure 21] (Linder & Ellis,
1990).
E African Mountain
South Africa
Figure 21: Worldwide Distribution of the species of Pentaschistis (Source: Google earth
Modified: www.google.com/earth/download/ 17th November 2010)
Pentaschistis (Nees) Spach, Hist. Nat. Veg. Phan. 13: 164 (1864); Stapf in F.C. 7: 480
(1899); McClean in S. Afr. J. Sci. 23: 273–282 (1926); Chippindal in Grasses & pastures, 254
103
(1955); Clayton in F.T.E.A. (Gramineae. Pt. 1): 124 (1970); Dyer, Gen. S. Afr. Pl. 2: 839
(1976); Clayton & renvoize, Gen. Gram. 174 (1986). Danthonia DC subgen. Pentaschistis
Nees in Linnaea 11: 125 (litt. ber.) (1837), Nees, Fl. Afr. Austr. 280 (1841). Type species:
Danthonia curvifolia Schrad.
Members of the genus Pentaschistis (Poaceae) are annuals or perennials with linear to folded
leaf blades. The inflorescences are panicles that are either open or contracted, occasionally
with glandular branches. The spikelets are 2−flowered, enclosed by the, lanceolate to oblong
glumes. These glumes are subequal, narrowly lanceolate to lanceolate, keeled, and 1−3
nerved. The lemmas are lanceolate to oblong, 5−9 nerved, 2−lobed, and awned from the sinus
of the lobes. The awns are geniculate, mostly with twisted column. The palea is narrow,
usually as long as the lemma. The callus is short and bearded (Linder & Ellis, 1990; Clayton,
1970; Phillips, 1995).
3.1.2 Nomenclatural background of the genus Pentaschistis
The first description of grasses that are currently included in Pentaschistis was by Thunberg
in his Prodromus Plantarum Capensium written and published on 1794 where he described
five species (two in Holcus L. and three in Avena L.). The next group of species were
described by Schrader, H.A. (1821) from specimens collected at Cape by Hesse where all of
them were placed in Danthonia DC. Nees ab Esenbeck (1832, 1841) recognized several
genera for the species now in Pentaschistis in Flora Capensis.
McClean (1926) transferred all the awnless two flowered species from Achneria to
Pentaschistis. He placed awnless species of Pentaschistis in Eriachne, while those with awns
in Danthonia section Pentaschistis. However, he placed P. capensis in the genus Triraphis.
Otto Stapf (1899, 1910, and 1915) raised the awned species of Danthonia section
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Pentaschistis to full genus Pentaschistis while retaining the awnless two flowered species in
the genus Achneria. He also placed Triraphis of Nees in Pentaschistis and the large spikelet
forms in Pentameris. Hubbard (1937) in Flora of Tropical Africa recognized 11 species of
Pentaschistis from Tropical African Mountains. He adopted a very narrow species concept
where he recognized a distinct species for each mountain block (Linder & Ellis, 1990). For
example, he recognized P. ruwenzoriensis C.E. Hubb. (from Mt. Ruwenzori), P. meruensis
C.E. Hubb. (from Mt. Meru), P. expansa (Pilg.) C.E. Hubb. (from Mt. Kenya) and P.
borussica from several mountains as distinct species. However all these four species, which
have open panicles, were merged under P. borussica (K. Schum.) Pilg. by Clayton (1970).
Clayton (1969) also transferred Eriachne malouinensis to Pentaschistis. Phillips, (1994)
reduced species with contracted panicles from various mountains previously treated as
distinct species (Clayton, 1970) in FTEA, namely P. minor and P. mannii as varieties under
P. pictigluma (Steud.) Pilg.
The researchers based their intuitive interpretations almost wholly on gross morphology and
were guided by inevitable differing intuitive taxonomic interpretations inherent in each
individual researcher, which consequently led to differing species concepts from the
narrowest (Linder &Ellis, 1990 and Clayton, 1970) to the broadest (Phillips, 1994 and 1995).
Hence Clayton, (1970) and Linder & Ellis (1990) recognized Pentaschistis pictigluma
(Steud.) Pilg. minor (Ballard & Hubbard) Ballard & Hubbard, P. gracilis S. M. Phillips, P.
mannii C.E. Hubbard, P. minor (Ballard & Hubbard) Ballard & Hubbard and P. imatogensis
C.E. Hubbard at specific rank while Phillips (1994,1995) place these taxa at infraspecific
ranks under the species P. pictigluma (Table 13) after failing to get clearly distinguishing
characters following her numerical analysis of several characters. This varying species
105
circumscription brings the total species recognized by various botanists to between six and
ten that occur in the tropical African Mountains.
Table 13: A summary of the species of Pentaschistis recognized by W. D. Clayton, S.
Phillips and P. Linder in East Africa and Ethiopian mountains
W.
D.
Clayton S. M. Phillips (1994 & H.P.
Linder
(1970)
1995)
Ellis( 1990)
P. chrysurus
−
P. chrysurus.
P. natalensis
−
P. natalensis
P. borussica
P. borussica
P. borussica
P. minor
P. pictigluma var. minor
P. minor
P. mannii
P. Pictigluma var. mannii
P. mannii
−
−
P. imatogensis
−
P. pictigluma var. pictigluma
P. pictigluma
−
P. pictigluma var. gracilis
P. gracilis
−
P. trisetoides
P. trisetoides
−
P. dolichochaete
−
&
R.P.
3.1.3 Previous phylogenetic analysis
Various molecular techniques are currently available for genetic diversity as well as
phylogenetic inferences including but not limited to restriction site analysis, comparative
sequencing, analysis of structural rearrangements and polymerase chain reaction (PCR) based
techniques (Baker et al., 1999). DNA sequencing techniques such as the one using nuclear
ribosomal DNA region and chloroplast DNA sequences are popular for phylogenetic studies
over a wide range of taxonomic levels. Recently, phylogenetic studies have focussed on the
use of chloroplast DNA (cpDNA) sequences because it is quite informative across many
taxonomic levels (Taberlet et al., 1991). The non-coding cpDNA region are more often used
for drawing phylogenetic relationships especially at lower taxonomic levels (Baker et al.,
106
1999) since they evolve more rapidly compared to the coding cpDNA regions thereby
accumulating insertion/deletion (Indels) at faster rate.
Phytogeographically, species of Afroalpine plants have complex derivation and are therefore
categorized into nine flora elements based on the supposed areas of origin of each species,
that is, the areas in which they have the majority of their closest relatives (Hedberg, 1961).
Despite the close proximity, the flora of the afro−alpine is floristically distinct and
ecologically isolated from the surrounding montane forest and savanna vegetation (Hedberg,
1970; Brühl, 1997; Menocal, 2004) and thus has been described by Hedberg (1970) as
isolated temperate “islands” since the low lying montane and savanna vegetation separate the
mountain enclaves from each other and from the rest of the floral elements. Hence the nearest
relatives of this Afroalpine enclaves are found not in the neighbouring tropical forest and
savannah but in alpine zones of Northern and Southern Hemispheres such as Eurasia (Boreal
element), the Mediterranian zone, Himalaya, Cape region and South Africa among others
(Hedberg, 1970). Therefore, like oceanic islands such as Galapagos and Hawaii, colonization
of this Afroalpine zone is thought for a long time to occur through infrequent long distance
dispersal from the nearest mainland or alpine zones (Hedberg, 1970; Lomolino et al., 2006).
Several recent phylogenetic studies using chloroplast DNA regions of common Afroalpine
genera such as Alchemilla, Arabis, Carex, Lychnis, Swertia and Ranunculus indicate that
species from these genera have colonized Afroalpine zones of tropical African mountains.
This happened through migration from distant areas such as Eurasia, Himalaya, South Africa
and Mediterranean areas among others (Koch et al., 2006; Assefa et al., 2007; Galley et al.,
2007; Popp et al., 2008; gehrke & Linder, 2009). Some recent phylogenetic and
phylogeographic studies combined the use of cpDNA and Internal Transcribed Spacer (ITS)
107
regions of nrDNA for phylogenetic reconstruction in the sub-family Danthonioideae (Baker
et al., 1995, 2000, 2003, 2007; Verboom et al., 2006; Galley & Linder, 2007; Pirie et al.,
2008) as well as improving the generic classification of this sub-family which is a
taxonomically difficult group (Linder & Ellis, 1990).
A species level phylogeny of the Pentaschistis clade (Pentaschistis, Prionanthium and
Pentameris) based on plastid DNA sequence data was constructed by Galley and Linder
(2007) to resolve phylogenetic relationships within the clade. In this study I attempted to
improve the phylogeny of Pentaschistis clade first by including all the species occurring in
the East Africa and Ethiopian Mountains absent from previous phylogenetic analysis as well
as using the ITS markers in addition to chloroplast sequence markers in resolving the
phylogenetic relationship of the Afroalpine sub–clade of the Pentaschistis clade.
Hence this study is giving the analysis, mainly morphological and genetic analysis that
hopefully will clarify the taxonomic treatment of Pentaschistis previously addressed by
researchers.
108
3.2 Materials and Methods
3.2.1 Sampling design and daw data collection
Herbarium voucher specimens were obtained from 75 plots in six mountains where plots of
100 × 100 m were sampled for morphological studies. For each population of Pentaschistis
occurring in the plots, 5 plants were sampled, one from each of the four corners and one from
the middle. Leaves from these plants were put in tubes containing silica gel for molecular
analysis and the first three plants were taken as vouchers for morphological reference and
verification. Identification was done both in the field and in the herbarium using literature
materials such as Floras e.g. FTEA, botanical books such as Agnew & Agnew, 1994, Ellis &
Linder, (1990). In addition to studying the species of Pentaschistis in the field, extensive
study of herbarium specimens were conducted at the East African herbarium at Nairobi,
Addis Ababa, Uganda, Tanzania and loaned materials from Kew herbarium including all type
specimens for detailed morphological study of species of Pentaschistis.
Tiny parts of the inflorescence such as the awns, glands, apical bristles and indumentums
were observed under dissecting microscope (De Wild.) and photos taken. Variations on
morphological characters such as culms length, lamina surface orientation, lamina width,
presence of glands on leaves, panicle shape, length, width and glands presence on panicles,
number of spikelets, spikelet length, glumes’ colour, apex shape, presence of glands, awn
length, apical bristle length and anther colour were recorded from at least five specimens in
each species from collected voucher specimens including previous herbarium collections and
data recorded in an excel spreadsheet for multivariate analyses (Cluster and Principal
Component Analysis).
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3.2.2 Sampling for phylogenetic analysis
Complete species sampling was attempted based on the taxonomy of Davidse (1988), Linder
& Ellis (1990), Galley & Linder (2006) and Phillips (1994; 1995). A total of 83 aligned
sequences (67 from gene bank, previous study and 16 from this study) representing 72
species (over 95% of species) were used in the phylogeny analysis. With the exception of five
Afroalpine species including the four varieties of P. pictigluma, the rest of the samples were
obtained from sequences of previous studies (Galley & Linder, 2007; Pirie et al., 2008)
deposited in the gene bank. Two Merxmuellera species were selected as outgroups. Most of
the samples used were silica dried although few herbarium specimens were used as well.
3.2.3 Phenetic analysis
Qualitative characters were coded and data matrix generated (Appendix 2a and 2b) for
phenetic analysis using STATISTICA version 9.0. Both cluster analysis and Principal
Component Analysis were conducted.
The distribution and habitat preference of the species of Pentaschistis were assessed from all
the sampled plots and from outside the plots. The cumulative frequency of occurrence of
Pentaschistis species were recorded on a table form and results presented in a simple line
graph.
3.2.4 DNA analysis
Total genomic DNA was extracted from silica dried specimens or herbarium sheet using the
standard CTAB protocol (Doyle & Doyle, 1987). Two non–coding regions of the chloroplast
DNA viz: TrnL-F and rpL-16 and internal transcribed spacer (ITS) from nuclear ribosomal
DNA were used for amplification and sequencing reactions. For both the plastid and the ITS
regions, PCR amplifications were performed in Thermocycler machine Gene Amp PCR
110
System 9700 (Applied Biosyst., USA) using a total reaction volume of 20 µl with 25mM
mgcl2, 10 × PCR buffer (Biolab, USA), 10 mM dNTPs, 10 µM primer and 1 unit of Taq
polymerase (Biolab, USA) and 2µl DNA template of unknown concentration. Table 14
shows the regions amplified and sequenced and the primers used.
Table 14: Gene region amplified and sequenced and the primers used for amplification
Gene Region
trnL-F
rpL16
Primers (PCR)
c, f
F71
R1000
Primers (sequencing)
c, d, e, f
F71
R1000
ITS
L/4
L/4
Source
Taberlet et al. 1991
Baum et al. 1998
Galley & Linder
2007
Baum et al. 1998
The trnL-F intron and intergenic spacer was amplified using a PCR protocol with an initial
denaturation step of 80°C for 5 minutes followed by actual denaturation at 94°C for 1 min,
followed by annealing temperature of 50°C for 1 min running for 35 cycles followed by an
extension of temperature at 72°C for 2 min and terminated with final extension of 5 min at 5
min. The rpL16 intron was amplified using an initial denaturation step of 94°C for 4 min
followed by 34 cycles of 1 min 94°C actual denaturation, 1 min at 55°C annealing step,
followed by 1 min at 72°C extension and a final extension of 7 min at 72°C.
The
amplification of the nuclear marker region of ITS followed PCR protocol with an initial
denaturation step of 4 min at 94°C followed by 35 cycles of actual denaturation of 30 s at
94°C, followed by annealing process for 1 min at 53°C, followed by initial extension for 3
min at 72°C and a final extension for 7 min at 72°C. PCR products were visualized on 2%
agarose gel for confirmation of successful amplification through band observation and
purified using the Agencourt AMPure XP system (Beckman Coulter, Brea, CA, USA) on a
Beckman Coulter Biomek (Beckman Coulter) robot using the manufacturer's protocol. Cycle
sequencing was performed on an Applied Biosystems 3730XL DNA Analyzer (Applied
111
Biosystems, Foster City, CA, USA) using the BigDye terminator version 3.1 with 5x
sequencing buffer (Applied Biosystems) following the manufacturer’s protocol.
3.2.5 Phylogenetic analysis
Bayesian analysis was performed on each of the markers separately and in combination using
Bayesian inference, as implemented in MrBayes 3.2 (Huelsenbeck & Ronquist, 2003).
Multiple sequence alignment was done using the ClustalW option in Bioedit version 7.0 and
adjusted manually. MEGA v5.05 (Tamura et al., 2011) was used to analyze the sequences’
characteristics, such as length of sequence, number of variable characters and parsimony–
informatives sites. The best fit substitution models for each of the three partions were chosen
using MrModeltest v2.3 (Posada & Crandall, 1998) as implemented in PAUP* v. 4.0b10
(Swofford, 2003) using the Akaike information Criterion (Akaike, 1973).
Bayesian analyses were run as two parallel MCMC (Marcov Chain Monte Carlo) analyses
using information from the model such as Nst, gamma/invigamma. For the separate analysis,
each run covered five and ten million generations for plastid and ITS respectively, with one
cold and three heated chains. Frequency sampling during the analysis for the plastid and ITS
regions occurred after every 5000 generation. The combined analysis covered 15 million
generation with sampling occurring after every 10, 000 generations. Convergence was
diagnosed using the standard deviation of split frequencies (SDSF) and the likelihood of each
parameter was checked for stability confirming sufficient convergence. In this case SDSF
value of between 0.007 and 0.009 was obtained for both the single gene and combined
analysis signaling enough convergences between the runs. Twenty five percent of the
generations were discarded as burn–in and remaining summarized in a 50%–majority rule
consensus tree with posterior probabilities (PP) as a measure of clade support.
112
3.3 RESULTS
3.3.1 Cluster analysis (CA)
A morphological data matrix involving 38 characters was analysed from 37 specimens
(OTUs=specimens) of Pentaschistis representing nine taxa. The results indicate three major
clusters. At linkage distance 100, two major clusters are visible that divides the tropical
Pentaschistis into two clusters. Cluster A comprises of specimens representing the two
narrow endemics, P. dolichochaeta and P. chrysurus in addition to the broadly distributed P.
natalensis. These are showing continuous variation.
Cluster B constitutes the broadly
distributed species of P. pictigluma variations (P. minor, P. mannii, P. pictigluma and P.
gracilis) and P. borussica. The only annual, P. trisetoides is nested within this cluster. At
linkage distance 60, three major clusters are evident and are: P. dolichochaeta, P. chrysurus
(Cluster A), P. natalensis (Cluster B) and P. mannii, P. minor, P. borussica, P. gracilis and
P. pictigluma (Cluster C).
160
140
120
100
80
60
40
113
P. pictigluma
P. pictigluma
P. pictigluma
P. pictigluma
P. borussica
P. borussica
P. borussica
P. borussica
P. borussica
P. gracilis
P. pictigluma
P. trisetoides
P. gracilis
P. mannii
P. mannii
P. mannii
P. mannii
P. mannii
P. minor
P. minor
P. minor
P. minor
P. minor
P. natalensis
P. natalensis
P. natalensis
P. natalensis
P. chrysurus
P. chrysurus
P. chrysurus
P. chrysurus
P. chrysurus
P. dolichochaeta
P. dolichochaeta
P. dolichochaeta
P. dolichochaeta
P. dolichochaeta
20
0
Figure 22: Phenogram of Custer Analysis for Pentaschistis species
Linkage Distance
114
3.3.2 Principal component analysis (PCA)
PCA is suitable for revealing patterns of continuous variation in a data set and allow us to
recognise the same groups defined by cluster analyses. Hence the same data matrix was
analysed using PCA except for one variable, leaf position that was not varying among the
taxa and was therefore eliminated from the analysis. The result of PCA of 37 morphological
characters showed similar pattern with that of Cluster Analyses (figure 23). Five groups are
visible: P. pictigluma variations, P. borussica, P. natalensis, P. chrysurus−P. dolichochaeta
and P. trisetoides. The first group on the top left are the two narrow endemic species P.
dolichochaeta and P. chrysurus. Below this group is P. natalensis which, in the study area
occur only in Tanzania. In the top right section of the PCA graph is the P. pictigluma species
complex comprising of the four varieties (P. pictigluma var. pictigluma, P. pictigluma var.
minor, P. pictigluma var. mannii and P. pictigluma var. gracilis). In the bottom right is the
other broadly distributed species P. borussica. In between P. pictigluma and P. borussica
occur the only annual P. trisetoides (Figure 23).
115
P. chrysurus-P. dolichochaeta
3
2
P. pictigluma species complex
1
PC2
0
-1
-2
-3
P. trisetoides
P. natalensis
-4
P. borussica
-5
-8
-6
-4
-2
0
2
4
6
PC1
Figure 23: Scatter plot of the PCA results of morphological variations for Pentaschistis species
116
Table 15: Distribution of Pentaschistis species in Eastern African mountains
Species
Pentaschistis
chrysurus
Altitude
(m)
Bale, Elgon, Kenya, All the four vegetation types 3159 to
Kilimanjaro, Rwenzori, but frequent in rock outcrop 4337
Abedares and Meru
and Alpine grasslands
Kilimanjaro and Meru Thicket/forest
2598 to
(Tanzania endemic)
3594
Pentaschistis
dolichochaeta
Ancober and Wafwasha Forest
forest
Pentaschistis
borussica
Pentaschistis
pictigluma
pictigluma
Pentaschistis
pictigluma
mannii
Pentaschistis
pictigluma
minor
Pentaschistis
pictigluma
gracilis
Pentaschistis
trisetoides
Mountain
Vegetation Community
2800 to
3000
Bale
and
Simen All vegetation types but 3875var. (Ethiopian Endemic)
frequent in rock outcrop and 4122
Alpine grasslands
Meru, Kilimanjaro and Grassland and rock outcrop
3900var. Kenya
3936
Bale, Kilimanjaro, Kenya All vegetation types, frequently 3636 to
var. and Elgon
in rock outcrops
4461
Bale
and
var. highlands
Bale and Simen
Ethiopian Grassland and Alchemilla
3705 to
4143
Rock outcrop
3705 to
3760
3.3.3 Distribution and Ecological Analysis of Pentaschistis Species across the vegetation
communities
Five species of the genus Pentaschistis (Poaceae) were sampled and recorded from the plots
and outside the plots (Table 15 and 16). Those sampled from the plots are three species
(figure 24) and include: P. borussica, P. pictigluma var. gracilis, P. pictigluma var. minor, P.
pictigluma var. pictigluma and P. trisetoides. Pentaschistis chrysurus and P. dolichochaeta
were sampled from outside the plots since they occurred below Afroalpine zone. Both species
usually occur in more or less forest zone or bush thickets near trails
117
Table 16: Frequency of Pentaschistis species in the five vegetation communities of
Afroalpine zone of Eastern Africa
Species
Pentaschistis borussica
Pentaschistis pictigluma var.gracilis
Pentaschistis pictigluma var. minor
Pentaschistis
pictigluma
var.
pictigluma
Pentaschistis trisetoides
Rock
outcrop
8
0
5
Bog
2
1
1
Grassland
7
0
2
Dendrosenecio
1
0
1
Alchemilla
0
1
0
5
2
4
0
2
0
0
0
0
0
Figure 24: Showing the occurrence and prevalence of five Pentaschistis species across
five vegetation types in East Africa and Ethiopian mountains
3.3.4 Molecular Analysis
The final matrix used for the phylogenetic analysis consisted of a total of 2032 plastid and
nuclear DNA sequence characters. Trimming at both ends of the dataset was done due to
118
some variations of some sequence length between the newly sequenced samples (current
study) and the retrieved sequences from the gene bank (previous studies). The lengths of the
sequences of trnL-F and rpL-16 regions are 693 bp and 767 bp respectively after the
trimming. For the aligned sequences of trnL-F the number of variable characters was
164(24.0%) of which 66 (9.5%) were parsimony-informative sites. A GTR+G substitution
model was selected for both partitions of the two plastids. The length of the ITS sequences
was 572 bp out of which were 180 (31.5%) variable sites and 93(16.3%) parsimony
informative sites. The best fit model for the ITS was GTR+ G + I.
From the Bayesian analysis, a consensus tree with fairly moderately resolved phylogenetic
tree (figure 25) is obtained which recovers most of the clades of the previous phylogenetic
analysis of Galley and Linder (2007). From this tree, the Afroalpine species from Eastern
Africa are grouped into two widely separated and distantly related clades that will form the
basis of the discussion below. The first Afroalpine species to form a clear sub-clade are the
two species P. borussica and P. pictigluma including all the four varieties of P. pictigluma
(P. pictigluma var. pictigluma, P. pictigluma var. minor, P. pictigluma var. gracilis and P.
pictigluma var. mannii).
Like in the previous phylogeny of Galley and Linder (2007) the sequence association or
grouping between these two species somewhat varied in that some accessions from one
species aggregate with another accession from the other species rather than an accession from
its own species (Sub-clade 1 figure 25). All the sequence accessions from Afroalpine species
are from this study but interestingly the same pattern is observed in the previous phylogenetic
analysis. The monophyly of both sub–clades are strongly supported.
119
Figure 25 Phylogenetic tree inferred from Bayesian analysis of the plastid DNA (trnL–F & rpL-16) and ITS data set of Pentaschistis
species. Posterior probability values are indicated above the branches
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3.4 DISCUSSION AND CONCLUSION
3.4.1 Phenetic and morphological analysis
The results from the morphological variation within and between the populations study
conducted in the Afroalpine zone of the Eastern Africa including morphometric analysis of
37 specimens for 38 characters of the species of Pentaschistis broadly indicates the existence
of two major groupings. The first of these groupings is the two widely distributed species
viz: P. borussica and the P. pictigluma species complex as well as two narrow endemics: P.
dolichochaeta and P. chrysurus. The two narrow endemic species are morphologically more
similar, as indicated by both the CA and the PCA. Study of herbarium specimens as well as
phenetic analysis (CA & PCA) also indicate that P. natalensis exhibit superficial
morphological similarity to the two vicariant narrow endemic taxa: P. chrysurus and P.
dolichochaeta that are endemic to Tanzania (Mt. Kilimanjaro and Meru) and Central
Ethiopian highland (Shewa region) respectively. Pentaschistis natalensis is the most widely
distributed species of the genus occurring from South Africa to southern Tanzania highland
and Madagascar. However, phylogenetic analysis of the Pentaschistis clade (Pentaschistis,
Pentameris Beauv. and Prionanthium Desvaux) based on the cpDNA characters by Galley &
Linder (2007) indicate that P. natalensis is not related to the tropical African Pentaschistis
and thus the morphological resemblance between P. natalensis and the two narrow endemics
is superficial.
Similarly comparative gross morphological study of P. pictigluma, P. minor, P. mannii and
P. gracilis indicate that there is overlap of characters among these taxa and hence are
phenetically not clearly distinct as revealed by both CA and PCA of the current study. About
19 characters are particularly important in separating well the taxa under study. These are:
culm length, leaf length and width, panicle length, number of spikelets, spikelet length, glume
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length & indumentums, anther length & colour, awn length, apical bristle length, lemma
length and indumentums, palea length, floret length and indumentums. However in the field,
subtle morphological differences between these taxa can be observed that distinguishes
between them. For example P. pictigluma var. pictigluma has characteristic golden yellow
panicles while P. pictigluma var. gracilis has silvery grey panicles and soft glandular leaves.
Pentaschistis pictigluma var. minor can be easily recognised by its short stature, few grey
spikelets and folded tough leaves.
The habitat preference and distribution of Pentaschistis species vary. Pentaschistis borussica
was the most widely distributed species as well as the most abundant species occurring in all
the mountains except Mt. Simen. This species frequently occur in rock outcrop and grass
vegetation communities but has also been recorded from bogs. It is rare in Dendrosenecio
and Alchemilla vegetation communities. Pentaschistis pictigluma var. minor was also widely
distributed occurring in four out of the six mountains sampled but was restricted largely on
rock outcrop vegetation community, most often in the mountain summits. Pentaschistis
pictigluma var. pictigluma, P. pictigluma var. gracilis and P. trisetoides are endemic to
Ethiopian Mountains including Bale and Simen. Pentaschistis pictigluma var. mannii. P.
chrysurus and P. dolichochaeta have been recorded from outside the plots. Pentaschistis
chrysurus is endemic to Tanzania while P. dolichochaeta is endemic to Shewa region of
Ethiopia.
Phenetically and morphologically there were significant differences between the broadly
distributed species (P. borussica–P. pictigluma) and the two narrow endemics (P. chrysurus
and P. dolichochaeta) thus rejecting the null hypothesis earlier stated in the hypothesis
section of the second chapter (hypothesis one). Habitat preference between these two
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groupings was also quite distinct as the narrow endemics are essentially forest (non-alpine
species) while the distribution range of the broadly distributed species was largely alpine
zone. Thus the two loose groupings exhibited altitudinal separation.
3.4.2 Phylogenetic analysis
A fairly moderately resolved phylogenetic tree of the Pentaschistis clade was obtained
revealing two colonization events of the Afroalpine species in East Africa and Ethiopian
Mountains. Consequently the earlier view held of single colonization event of tropical
African Pentaschistis is rejected (null hypothesis two of the second chapter). The
Pentaschistis clade consists of three genera viz: Pentameris, Pentaschistis and Prionanthium,
and was so named by Barker et al. (2000) following the phylogenetic analysis of the DNA of
Danthonioideae grasses. The groupings of these three genera are supported by morphological
characters such as spikelet similarities-presence of multicellular glands, having two fertile
florets per spikelet and rachilla extension. They also share weak development (loss of)
haustorial synergids, fine granular or lack of starch in the synergids compared to the presence
of globular starch in other Danthonioid genera, insertion of the lemma setae in the sinuses
between the lateral lobes and the median awn. In addition, these three genera have basic
chromosome number of x= 7.
This morphological and cytological evidence is strengthened by the recent phylogenetic
analysis of Galley and Linder (2007) as well as this study that showed the two genera of
Pentameris and Prionanthium are nested within the Pentaschistis clade. However, since the
scope of this study is limited to tropical African Mountains, this discussion will focus on the
Afroalpine sub-clades of the genus Pentaschistis that constitute the bulk of the species of the
Pentaschistis clade. Although most of the clades described in Galley & Linder (2007) are
123
recovered in the resulting phylogenetic tree, the discussion will only focus on the clades
nesting the Afroalpine sub-clades.
As revealed by the phylogenetic tree, the Afroalpine species of Pentaschistis grouped into
two distantly related clades with high support of posterior probability (PP= 100). This
provides strong evidence of two independent events of colonization of Pentaschistis species
in the Afroalpine zones of tropical East Africa and Ethiopian Mountains from the source
region (Cape Floristic Region and South Africa). The first Afroalpine species sub–clade are
grouped with the large “summer rainfall clade” (VII). Basically this first group consists of the
two broadly distributed species: P. borussica and P. pictigluma and the only annual species
P. trisetoides that is sister to the P. pictigluma–P. borussica sub-clade. Other members of the
“summer rainfall clade” apart from the Afroalpine species are: P. lima, P. aristifolia, P.
tomentella, P. airoides ssp. airoides, P. pseudopallescens, P. veneta, P. tomentella, P.
oreodoxa, P. glandulosa, P. natalensis, and P. insularis among others. All the variations of P.
pictigluma (var. pictigluma, var. minor, var. mannii and var. gracilis) are all represented as
well as the different collections of P. borussica from East Africa and Ethiopian Mountains.
Both the morphological and morphometric analysis of the current study as well as previous
morphological variation study indicate almost lack of distinct morphological characters that
separates the P. pictigluma varieties and to a lesser extent between P. pictigluma varieties and
P. borussica. Similar trends where sequences from P. borussica and P. pictigluma are
grouped together in the internal nodes are observed. From these observed trends and the wide
distribution of these two species, it can therefore be inferred that these (P. borussica and P.
pictigluma) are young taxa undergoing rapid diversification following their introduction in
the Afroalpine zone as well as lineage sorting.
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The second sub-clade of Afroalpine species includes the two narrow endemics viz: P.
chrysurus and P. dolichochaeta that aggregate with P. basutorum and P. juncifolia which
form a grade with strong support (PP= 99; sub-clade 2). Pentaschistis chrysurus and P.
dolichochaeta share a number of morphological characteristics such as robustness (size),
relatively broad leaves, long culms, large spikelets and florets, long awns and lack glands.
These two species occur in similar habitats (forest or thicket) and occupy between forest zone
and sub-alpine zone (2500m to 3400 m). Phillips (1995) in Flora Ethiopia and Eritrea
similarly noted the close morphological affinities between these species. Both Pentaschistis
dolichochaeta and P. chrysurus are endemic to Ethiopia and Tanzania respectively where the
former is limited to Shewa region specifically in Ancober and Wafwasha natural forest while
the latter is limited to Mt. Kilimanjaro, Mt. Meru and has been cited from Mt. Hannang all in
northern Tanzania (Phillips, 1995).
Generally speaking species after their introduction to new suitable habitats tend to spread,
diversify and occupy as large area as possible depending on the success of any given species
before it starts shrinking its range due to natural or anthropogenic factors, natural
evolutionary process that ultimately lead to extinction. Since rarity precedes extinction, the
species continue to decline in terms of population density and geographic range until it
ultimately disappears altogether (becomes extinct). From biogeographical point of view
therefore, these two species are undergoing range shrinkage where their habitat is
diminishing implying they are old taxa on road to extinction compared to P. pictigluma and
P. borussica that are undergoing rapid diversification and colonizing available Afroalpine
habitats in East African and Ethiopian Mountains. Hence these two species are perhaps
vulnerable to extinction since their habitats (forest/thickets) in the lower zones of the high
125
mountains are experiencing anthropogenic induced destructions and fragmentation that is
causing loss of biodiversity.
The current phenetic study, gross morphology and DNA sequences of tropical African
Pentaschistis reveal the distinctness of P. chrysurus, P. dolichochaeta from the rest of the
taxa and the close morphological affinities between them. Pentaschistis borussica can be
easily distinguished from the four varieties of P. pictigluma by its raised glands and is also
phenetically distinct from these taxa. The phenetic analysis results (and sequence similarities)
among P. pictigluma, P. minor, P. gracilis and P. minor indicate that these taxa are not
distinct from each other. Hence this study proposes the retention of the current taxonomic
treatment. It also further recommends population genetic studies among the populations of P.
pictigluma species complex and between the two narrow endemics, P. dolichochaeta and P.
chrysurus to reveal the existence of any cryptic speciation and possible hybridization.
3.4.3 Key to Pentaschistis species occurring in African high mountains
This section gives dichotomous key that shows the species circumscription adopted by this
study after the various analysis.
1. Plant annual; lacking rhizomes/stolons……………….....................Pentaschistis trisetoides
Plant perennial; rhizomes/stolons mostly present…………………………..........................2
2. Inflorescence loose and/or open; 3–17 cm wide……………………………........................4
Inflorescence loosely contracted to linear/spiciform; 0.8–2.5 cm wide……........................3
3. Culm length 60–120 cm; lamina length 40−60 cm; spikelet length 8–9 mm; endemic to
Shewa region of Ethiopia………………………………......................P. dolichochaeta
Culms length 6−40 cm; Lamina length 3−20 cm; spikelet length 4–7 mm long, widespread
in tropical African
Mountains…………………….................................P. pictigluma
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4. Leaf margins and inflorescence glandular; florets glabrous, 2.5–3 mm; widespread in East
Africa and Ethiopia…………………………………………………..........P. borussica
Leaf margins and inflorescence eglandular; florets densely hairy, 4 mm; restricted to
Tanzania…………………………………………………….........................................5
5. Panicle ovate to oblong, florets not intertwined; spikelet number over 200; glume apex
acuminate…………………………………………….................................P. chrysurus
Panicle pyramidal in shape, florets intertwined; spikelets 35–150; glume apex
acute…………………………………………………….............................P. natalensis
Varieties of P. pictigluma
a)
Golden brown panicle; lamina flat or in−rolled, may or may not be glandular
…………………………………......................................................................... var. pictigluma
b)
Grey or silvery panicles, not glandular; lamina linear, flat, margins usually
glandular…………………………............................................................................var. gracilis
c)
Eglandular plants; panicles grey few spiculate; lamina narrowly linear or in−rolled to
setaceous………………………..................................................................................var. minor
d)
Panicles grey, occasionally glands restricted to pedicels, raised crateriform, leaf
sheaths white, shiny & persistent; …………………….............................................var. mannii
1. Pentaschistis trisetoides (Hochst. ex Steud.) Pilg. In Notizbl. Bot. Gt. Berlin 9:516
(1926); C.E. Hubb., F.T.A. 10: 131 (1937); Cufodontis, Enum. Ethiop. 1235 (1968); Linder
& Ellis, Contrib. Bol. Herb. 12:50 (1990); S.M. Phillips, F.E. & E. (1995); Type: Ethiopia,
near Debra Eski, Schimper 109 (K, holo!, NH, ETH, iso!).
Synonym: Danthonia trisetoides Hochst. ex Steud., Pl. Glum. 1:244 (1854); K. Schum.,
Engl., Pflanzenw. Ost−Afrika C: 109 (1895). Danthonia segetalis Hochst. in Flora 38:276
(1855). Pentaschistis segetalis (Hochst.) Pilg. In Notizbl. Bot. Gt. Berlin 9:518 (1926). Type:
127
Ethiopia, near Debra Eski, Schimper 24 (TUB, holo., K). Danthonia trisetoides Hochst. ex
Steud. var. tenuis Engl., Hochgebirgsflora 131 (1892). Pentaschistis trisetoides (Hochst. ex
Steud.) Pilg. var. tenuis (Engl.) Pilg. In Notizbl. Bot. Gt. Berlin 9:517 (1926). Type: Ethiopia,
Schimper 621 (B, holo., K), Schimper 766.
Slender annual glandular plant with simple culm, 6–35 cm tall. Leaf lamina soft linear, flat,
glabrous above, pubescent below, 2–10 cm × 0.2 cm wide, margin scabrid. Inflorescence
open or loosely contracted. Panicles glandular with raised, crateriform glands. Spikelets about
45–100, silvery grey with purple tinged, 3.5–5 mm long. Glumes lanceolate, acuminate
greenish purple 3.5–5 mm long glabrous to scaberulous occasionally glandular. Florets
glabrous, 2 mm long. Lemma pubescent, 1.5–2 mm long, awned and with two apical bristles.
Apical bristles 1–2 mm long. Palea glabrous, 1.5−2 mm long. Awn 5–6.5 mm long,
geniculate. Anthers brown 0.3–0.7 mm long.
Voucher specimen(s) cited: Ethiopia, Gondar region, Mt. Simen, Geech, AFROALP II team,
ET 220−3, 25 October 2007.
Distribution: Bale, Simen & Ethiopian highlands (TU, BA, GD, SU, AR)
2. Pentaschistis dolichochaeta S.M. Phillips; Type: Ethiopia, SU, Ankober, Lemma
Selassie 887 (Eth holo.).
Perennial eglandular fairly robust plant with rhizomatous base. Culm 60–120 cm tall. Lamina
40–60 cm × 0.3−0.6 cm, flat occasionally inrolled 3–6 mm wide, margins scabrid, puberulous
on the upper surface glabrous below. Panicle open to loosely contracted 10–16cm × 3–4 cm.
Spikelets 180–200, pale greenish yellow 8–9.5 mm long. Glumes pale green, acuminate,
glabrous or nearly so 8–9 mm long. Florets densely hairy, 4 mm long. Lemmas 3–3.5 mm
long, pilose with two apical bristle 5–6 mm long arising from the lemma lobes, palea
128
glabrous, 4−5 mm long. Awn 9–15 mm long, geniculate, column 1–2 mm long. Anther
golden yellow, 2.5–2.7 mm long.
Ethiopia, Shewa region, Debre Sina, Wafwasha natural forest, 24 April 2009, Ahmed A. Abdi
ET−1604−1, ET−1604−6, ET−1604−7, ET− 1604−8, 1 ET−604−21.
Distribution: Wafwasha forest and Ancober, Shewa (Ethiopia, SU). Not known elsewhere;
2920−3000 m.
3. Pentaschistis borussica (K. Schum.) Pilg. In Notizbl. Bot. Gt. Berlin 9: 517 (1926); Hubb.,
F.T.A. 10:128 (1937); Clayton, F.T.E.A. 1: 125 (1970); Linder & Ellis, Contrib. Bol. Herb.
12:55−56 (1990); Danthonia borussica K. Schum. In Eng., Pflanzenw. Ost−Afr. C: 109
(1895). Type: Tanzania, Mt. Kilimanjaro, Volkens 1368 (EA. iso!, B, holo., BM, K).
Synonym: Pentaschistis trisetoides (Hochst. ex Steud.) Pilg. var. expansa Pilg. in Notizbl.
Bot. Gt. Berlin 9: 516 (1926).
Pentaschistis expansa (Pilg.) C.E. Hubb. in F.T.A. 10:130 (1937). Type: Kenya, Mt. Kenya,
Fries 1200b (B, holo.).
Pentaschistis effusa Peter in FDOA 1, Anh.: 97 t. 56/1 (1930). Type: Tanzania, Mt.
Kilimanjaro, Peter 46685 (B, holo.).
Pentaschistis meruensis C.E. Hubb. In Kew Bull. 1936:501 (1936); C.E. Hubb., F.T.A
10:127 (1937). Type: Tanzania, Arusha District, Mt. Meru, Burtt 4062 (K, holo!, EA, iso!).
Pentaschistis ruwenzoriensis C.E. Hubb. In Kew Bull. 1936:500 (1936); C.E. Hubb., F.T.A
10:127 (1937). Type: Uganda, Toro District, Mt. Ruwenzori, Taylor 2903 (K, holo., EA iso!)
129
Tufted perennial glandular plant with short rhizome. Culm, 11–60 cm tall. Leaf lamina
linear, flat or folded, pubescent above, glabrous below 5–25 cm × 0.2−0.4 cm, margin
smooth, sheath mouth villous. Inflorescences open with 20−70 spikelets. Panicles glandular
with raised glands on the pedicels, branches, glumes and awns, 3−16 × 1−15 cm. Spikelets
light to dark grey , (3.5)4–6(8) mm long. Glumes lanceolate−oblong, acute or acuminate
greenish to pale yellow often purplish tinged 4–6 mm long glabrous to scaberulous
occasionally glandular. Florets glabrous, 2.5−3 mm long. Lemma glabrous or pubescent, 2–3
mm long, awned and with two apical bristles. Awn geniculate, 5–9(11) mm long, apical
bristles 2–3 mm long. Palea glabrous, 2−4 mm long. Anthers golden brown to purple 0.9–2.5
mm long.
Tanzania, Moshi, Arusha National Park, Mt Kilimanjaro, Shira, 6 November 2008, Abdi, A.,
Abel G., Wondimu T., C. Masao & FellyT., TZ−116−4; 10 November 2008, Karanga,
TZ−184−4; Kenya, Kitale, Mt. Elgon, east of Koitobos, 27 January 2009, Abdi et al.
KN−251−3, KN−251−4; Nyeri, Aberdares, Satima peak, 12 February 2009, Abdi et al.
KN−538−3.
Distribution: Bale and southern highland, Ethiopia (BA, AR, SU), Mt. Kenya, Elgon,
Aberdares, Kilimanjaro, Meru & Ruwenzori (U1−3, K3, 4; T2); 3000−4680 m.
3.
Pentaschistis chrysurus (K. Schum.) Peter in Fedde Repert. Beih. 40.1:303(1931);
Hubb., F.T.A. 10:125 (1937); Clayton, F.T.E.A. 1: 124 (1970); Linder & Ellis, Contrib. Bol.
Herb. 12:105(1990).
Synonym: Danthonia chrysurus K. Schum. In Eng., Pflanzenw. Ost−Afr. C: 110 (1895).
Type: Tanzania, Mt. Kilimanjaro, Volkens 1826a (B, holo., EA. iso!,K, photo).
130
Robust perennial eglandular plant with rhizome. Culm, 84−150 cm tall. Leaf lamina tough,
linear, flat, glabrous above, pubescent below 43–56 cm × 0.7−1.0 cm, margin smooth and/or
folded, sheath mouth villous. Inflorescences open with 200−250 spikelets. Panicles open,
glabrous, dense, much branched,10−32 × 3−8 cm. Spikelets light to dark brown,7.5–10 mm
long, ultimate pedicels (1)4−8 mm. Glumes acuminate light yellow to golden yellow, purple
tinged, glabrous 7.5–10 mm long. Florets pubescent, 4 mm long. Lemmas pubescent, 3–5
mm long, awned and with two apical bristles. Awn geniculate, 8–15 mm long, apical bristles
3–6 mm long, column1−2 mm, slightly twisted . Palea glabrous, 4.5−5 mm long. Anthers
golden brown 2–2.5 mm long.
Distribution: Mt. Kilimanjaro, Meru & Hanang (Tanzania, T2); 2598−3594 m.Tanzania, T2,
Moshi, Mt. Kilimanjaro, 19 November 2008, Abdi, A., Abel G., Wondimu, T., Masao, C. &
Felly, T. TZ−348−4; Mt. Kilimanjaro, 26 February 1934, Greenway 3801; 2 December 1972,
Vesey−FitzGerald D. 7506; Arusha National Park, Mt. Meru, 24 September 1967, Gilbert,
V.C. 2249; Meru crater, 28 December 1965, Vesey−FitzGerald D. 4853.
4.
Pentaschistis pictigluma (Steud.) Pilg. In Notizbl. Bot. Gt. Berlin 9:517 (1926); C.E.
Hubb., F.T.A. 10: 133 (1937); Cufodontis, Enum. Ethiop. 12343 (1968); S.M. Phillips, F.E.
&E. 7:70−71 (1995); Linder & Ellis, Contrib. Bol. Herb. 12:63−64 (1990).
Synonym: Aira pictigluma Steud., Pl. Glum. 1: 221 (1854). Type: Ethiopia, Schimper;
Danthonia anothoxanthiformis Hochst. (1855) nom. Superfl.−Type: Ethiopia without precise
locality, Schimper s.n. (P, holo.).
Danthonia thumbergii sensu A. Rich. Non Kunth; based on Quartin Dillon & Petit s.n. (P);
Danthonia uberior Hotchst. In Flora 38:279 (1855). Types: Ethiopia, Simien, mt. Bachit,
Scimper 112 (K, P); Schimper 541 (K);
131
Danthonia nana Engl., Hochgebirgsflora 131 (1892). Type: Ethiopia, on Mt. Guna, Schimper
1561 (B, holo., BM, K);
Danthonia trisetoides Hochst. ex Steud. var. schimperi Engl., Hochgebirgsflora 130 (1892).
Pentaschistis trisetoides (Hochst. ex Steud.) Pilg. var. Schimperi (Engl.) Pilg. In Notizbl. Bot.
Gt. Berlin 9:517(1926). Type: Ethiopia, Amba Hedscha, Schimper 1005(B, holo.,K).
Tufted very variable perennial plant with or without glands with short rhizome. Culm, 7−50
cm tall. Leaf lamina filiform and inrolled to soft and flat, glabrous or pilose 4–20 cm ×
0.2−0.4 cm, margin scabrid, sheath mouth villous, basal leaf sheath whitish and papery.
Inflorescences contracted or loosely contracted to spiciform with 10−100 spikelets. Panicles
elliptic to narrowly oblong, glabrous to puberulous, 2−13 × 0.5−0.8 cm. Spikelets golden
brown or grey to silvery grey,4–7 mm long, ultimate pedicels 2−3 mm. Glumes obtuse or
acute or acuminate grey green to pale yellow, purple tinged, glabrous to scaberulous 4–7 mm
long. Florets pubescent, 2−3 mm long. Lemmas pubescent, 2–2.5 mm long, awned and with
two apical bristles. Awn geniculate, 3.0–7 mm long, apical bristles (1)1.5–3 mm long,
column1−3 mm, twisted . Palea glabrous, 2−3 mm long. Anthers golden brown or yellow
brown, 0.3–1.2 mm long.
(a) P. pictigluma var. pictigluma
Characteristic golden brown panicles; spikelets (4.5) 5.5−7(7.7) mm long; awns (4) 5−7 mm
long; leaf blades flat or more often inrolled.
Ethiopia, Bale, Bale Mountain National Park, Sannati plateau, Konte, 19 November 2007,
AFROALP II Team ET−902−3; 12 November, AFROALP II Team ET−647−3; Garba
Guracha, 15 November 2007, AFROALP II Team ET−763−3; Angasu, 20 November 2007,
132
AFROALP II Team ET−813−3; Gondar region, Simen, Simen Mountain National Park,
Chenek, 28 October 2007, AFROALP II Team ET−366−1.
Distribution: Bale, Simen and Ethiopian highlands above 3500 m(Ethiopia, GD, BA, AR,
SU, GJ,TU); 3700−4200 m.
(b) P. pictigluma var. gracilis (S.M. Phillips) S.M. Phillips in Proc. 13th AETFAT Congress:
372 (1994); Synonym: P. gracilis S.M. Phillips in Kew Bull. 41: 1028(1986); Linder & Ellis,
Contrib. Bol. Herb. 12:64−65 (1990); Type: Ethiopia, SU, Entoto hill, Friis et al. 1303 (K,
holo.).
Silvery grey panicles; spikelets(3.5)4−6 mm long; awns 3−5 mm long; leaf blade linear, flat
and soft, margins usually glandular.
Ethiopia, Bale, Bale Mountain National Park, Sannati plateau, Angasu, 23 November 2007,
AFROALP II Team ET−1009−3; Garba Guracha, 15 November 2007, AFROALP II Team
ET−717−1.
Distribution: Mt Bale and South & Central highlands (Ethiopia, BA, SU, KF); Elgon
(Kenya/Uganda, T3).
(c) P. pictigluma var. minor (Ballard & C.E. Hubb.) S.M. Phillips in Proc. 13th AETFAT
Congress:371 (1994); Synonym: P. borussica (K. Schum.) Pilg. var. minor Ballard & C.E.
Hubb. In Kew Bull. 1930: 121 (1930); P. minor (Ballard & C.E. Hubb.) Ballard & C.E.
Hubb. In F.T.A. 10: 132 (1937); Cufodontis, Conspec. Fl.Aeth. 1234 (1968); Clayton,
F.T.E.A. 1:127 (1970); Linder & Ellis, Contrib. Bol. Herb. 12:62 (1990); Type: Tanzania,
Mt. Kilimanjaro, Cotton & Hitchcock 64 (K, holo.!).
133
Eglandular plants, comparatively short perennial variety; panicles grey few spiculate;
spikelets 3.6−5 (6) mm long; awn 2.5−5 (6) mm; leaf blade narrowly linear, folded to
setaceous.
Kenya, Kitale, K5, Mt. Elgon, eastern slopes, 3 March 1956, Bogdan, A. AB 4123; Upper
heath zone, 1 January 1957, Bogdan, A. AB 4499; Mt. Elgon, December 1933, Dale, I.R.
R3189; Ethiopia, Bale, Bale Mountain National Park, Sannati plateau, 12 November 2007,
AFROALP II Team ET−946−3; Tanzania, T2, Moshi, Mt. Kilimanjaro, Shira,4 November
2008, Abdi, A., Abel G., Wondimu, T., Masao, C. & Felly, T. TZ−48−4.
Distribution: Bale (Ethiopia, TU, AR & BA); Mt. Kenya, Elgon (Kenya &Uganda, K5, 4,
U3); Kilimanjaro (Tanzania, T2); 3000−4800 m.
(d) P. pictigluma var. mannii (Stapf ex C.E. Hubb.) S.M. Phillips in Kew Bull. 1936: 501
(1936); C.E. Hubb. In F.T.W.A. 2: 528 (1936); F.T.A. 10: 134 (1937); Clayton, F.T.E.A.
1:127 (1970); Linder & Ellis, Contrib. Bol. Herb. 12:62 (1990); Type: Cameroun, Mt.
Cameroun, Mann 1351 (K, holo.!)
Panicles grey, occasionally glands restricted to pedicels; spikelets 6−8 mm long; awn 5−7
mm long; leaf blade flat or folded, leaf sheaths white, shiny & persistent.
Tanzania, T2, Arusha National Park, Mt. Meru, 28 November 2008, Gilbert, Abdi et al.
TZ−453−4; TZ−452−4; TZ−454−5; western slopes above Olkakola estate, 27 October 1948,
Hedberg, O. 2301; Cameroun, Mt. Cameroun, 10 April 1955, Thresh, J.M. JMT 4.
Distribution: Mt. Elgon, Moruongole (Uganda, U1, 3); Meru (Tanzania, T2); Mt. Cameroun
(Cameroun); 2600−4500 m.
134
CHAPTER FOUR: SUMMARY DISCUSSION, CONCLUSIONS AND
RECOMMENDATIONS
4.1 Summary discussion
Afroalpine “Sky Islands” of tropical Eastern Africa forms an excellent natural laboratory to
study the biodiversity dynamics and the influence of abiotic and possibly biotic factors that
ultimately cause diversification of Afroalpine species following successful dispersal from the
source region. A total of 46 vascular plant families consisting of 124 genera and 278 species
were recorded and sampled from five vegetation communities rock outcrops, bogs, grassland,
Dendrosenecio forest or woodland and Alchemilla across six high mountains viz: Mt Kenya,
Elgon, Kilimanjaro and Ruwenzori, Bale and Simen. Interestingly, the result indicates the
dominance of few families over the majority in the colonization and occurrence of their
species and genera in the sampled vegetation types and Mountains while 65% of the
Afroalpine families have monotypic genera that are mostly represented by one or two species.
For instance Asteraceae and Poaceae, the two dominant families have 21 and 17 genera
respectively and account for more than a third (38%) of all the species of Afroalpine
collected from the sampled plots while families like Rubiaceae, Asphodolaceae, Juncaceae,
Geraniaceae, Commelinaceae and Callitrichaceae among others (30 families) are each
represented by only one genus that has one or less often two species.
A number of factors could have contributed to the success of Asteraceae and Poaceae such as
morphological adaptations including development of tiny light seeds especially grasses and
parachute structures like most of the members of Asteraceae that enable the species to be well
adapted for long distant dispersal by wind. This is because wind dispersal is thought to be one
of the main mechanisms effecting long distance dispersal for Afroalpine plants (Troll, 1952;
Hedberg, 1964b &1970; Coe, 1967) since all the Afroalpine enclaves are effectively isolated
from other flora of the surrounding vegetation and from each other by the high elevation. In
135
addition, wind dispersal or anemochory is characteristic of pioneer vegetation and virgin
habitats like newly formed volcanoes or after receding of glaciers or oceanic islands (Van der
Pijl, 1969) and its significance is directly proportional to the altitudinal height of the
mountains (Hedberg, 1970). Strong winds and occasionally wind storms like cyclones are
prevalent in Afroalpine zones of the high mountains and are capable of blowing large amount
of light and tiny seeds between the mountain Afroalpine enclaves and even from far distant
areas in the Northern and Southern Hemispheres with alpine climate. Other factors such as
fire (especially favours grass community), physiological and genetic adaptations could be
playing role in the dominance and rapid diversification of some families over others but is out
of scope of the current study (Wesche et al., 2000).
The Afroalpine plant communities prevalent in the five vegetation types sampled were
dominated by one or few keystone genera and species. The following are the main
combination of the keystone genera and species that best describes the communities in the
vegetation type: Carex runssoroensis−C. monostachya−Deschampsia caespitosa dominated
bogs, Festuca abyssinica−F. simensis−Pentaschistis dominated grassland communities,
Dendrosenecio−Alchemilla species dominated Dendrosenecio forest/woodland, Alchemilla
argyrophylla−A. johnstonii−A. haumannii dominated shrubby Alchemilla communities and
Helichrysum−Festuca−Koeleria−Pentaschistis dominated rock outcrop communities.
In the rock outcrop vegetation type, the species combination of the dominant genera like
Helichrysum, Festuca, Pentaschistis and Deschampsia generally varies from mountain to
mountain but the trend was that East Africa Mountains like Mt. Kenya, Elgon and
Kilimanjaro (all on the eastern arm of the rift valley) share common species combination
compared to the Ethiopian Mountains or those mountains occurring along the western arm of
136
the rift valley. Some genera in the rock outcrop were mountain specific and include
Rytidosperma (R. subulata), Colpodium (C. hedbergii), Plantago (P. afra), Cotula (C.
abyssinica), Gnaphalium unionis, Vulpia bromoides, Crepis rueppellii and Lychnis
abyssinica in Simen; Cineraria deltoidea, Crassula sp. in Bale; Dendrosenecio keniodendron
in Mt Kenya and Isolepis sp. in Kilimanjaro.
Unlike in the rock outcrop communities, bog vegetation types were more homogenous in the
generic composition. Mount Rwenzori, the wettest mountain differs from the rest of the
mountains with respect to species composition of the most dominant taxa. For example the
tussock forming species of Carex runsoroensis was the dominant species in all the four plots
of bog sampled from Ruwenzori while the rest of the mountains were dominated by Carex
monostachya in association with various members of other genera like Ranunculus (mostly R.
oreophytus), Haplocarpha (usually H. rueppellii) and Isolepis among others. Frequently
present in the tussocks of C. runnsoroensis are Hydrocotylee sibthorpioides, Isolepis fluitans,
Helichrysum
stuhlmannii,
H.
formosissimum
and
Alchemilla
argyrophylla
ssp.
argyrophylloides.
The genera and species of the family Poaceae form the dominant species in terms of cover in
grassland community. Species of Festuca (F. abyssinica in EA Mountains and F. simensis in
Ethiopian Mountains) dominated the species composition of the sampled plots across the
mountains. Other dominant grass genera in East Africa Mountains include Koeleria, (K.
capensis) Pennisetum (P. humile), Colpodium, Pentaschistis (P. borussica) and Poa. In
Ethiopian Mountains, Festuca (F. simensis), Pentaschistis (P. pictigluma), Andropogon (A.
lima) and Helictotrichon (H. elegantum) dominated grassland species
137
The Dendrosenecio is not found in Ethiopia, as a result only three East Africa countries viz.,
Mt Kenya, Elgon (Kenya/Uganda) and Ruwenzori were covered. However in Mt
Kilimanjaro, the Dendrosenecio vegetation type was not well developed as only scattered
trees were found in the Afroalpine zone which could not form continuous plot of
Dendrosenecio community comparable to other mountains for sampling and analysis. Mount
Kenya, Elgon and Ruwenzori each had an endemic vicariant species of Dendrosenecio as the
dominant species associated frequently and/or co−dominated occasionally with other woody
(Lobelia) or shrubby Alchemilla species. For example, in Mt Elgon D. elgonensis was the
dominant species while D. keniodendron dominates Mt Kenya Dendrosenecio woodland. In
Ruwenzori, Dendrosenecio vegetation type was well developed often forming dense moist
forest dominated by D. advinalis ssp. advinalis and D. advinalis var. petiolatus.
Alchemilla community was the least extensively developed vegetation type in the whole of
the study area although well represented by many species across the entire mountains. As
such only eight plots were sampled from two mountains (Bale in Ethiopia and Ruwenzori in
East Africa) and the other mountains lacked sizable plots for sampling and analysis. In
Ruwenzori the Alchemilla communities were dominated in terms of species cover and
presence by A. argyrophylla ssp. argyrophylloides and A. johnstonii with frequent association
of A. triphylla. On the other hand, A. haumanii was basically the sole dominant species in
Bale often with cover of over 84%. In one plot however, A. micrbetula, A. pedata and
Isolepis fluitans (all frequently found in bogs) dominated the Alchemilla community.
Rock outcrop and grassland vegetation types were the most similar (J’ value of 52) sharing a
total of 118 species. The other pairs that share relatively higher degree of similarity were
bogs and grassland (J’=44.0; common−96; different 122) and rock outcrop and bog (J=41.0;
138
common−101; different−143). On the other hand there were marked dissimilarity between
grassland and Dendrosenecio vegetation types (J=22.0; common−40; different−145) and/or
rock outcrop and Alchemilla (J=26.0; common−55; different−145). Light, water and
temperature play significant role in the occurrence and distribution of species in the
vegetation types. Shade loving species frequently occur in Dendrosenecio and Alchemilla
vegetation types since the dense forest or sclerophyllous shrubby Alchemilla provide cover to
the undergrowth species.
Analysis of similarities (R value) reveals the differences between the five vegetation types
i.e. can be easily recognized from each other. However the heterogeneity of the vegetation
types was blurred by overlap in terms of species composition. For instance vegetation types
such as rock outcrop vs. grassland were barely separable at all (R< 0.25) while rock outcrop
vs Dendrosenecio, rock outcrop vs Alchemilla, bog vs Alchemilla, bog vs grassland,
grassland vs Alchemilla, and Dendrosenecio vs Alchemilla were overlapping but fairly
separable. However, rock outcrop vs bog and bog vs Dendrosenecio are overlapping but
clearly separable (R > 0.5).
The rock outcrop vegetation type was the most species rich compared to both Dendrosenecio
and Alchemilla vegetation types. This was also true between bogs and Dendrosenecio
vegetation types where single factor ANOVA tests gives a marked differences between these
two vegetation types.
Comparative analysis of the species composition and community ecological analysis revealed
a general pattern of similarity and/or differences between the six mountains based on several
parameters such as geographical location, age, geological history (nature of formation),
139
climatic and edaphic or soil factors. Consequently East Africa Mountains that occur along
eastern arm of the rift valley such Mt. Kenya, Kilimanjaro and Elgon have high degree of
similarity compared to Ethiopian Mountains and Ruwenzori. Hence Mt Kenya, Kilimanjaro,
and Elgon share many common species and are floristically similar. Mount Kenya and
Kilimanjaro, the two closest mountains in East Africa show the high degree of similarity
(Jaccard’s similarity index: 44.4). Similarly Bale and Simen have higher Jaccard’s similarity
index than between any other pair for the same reason (J’ 47.05). Apart from geographical
proximity, age, nature of formation and climatic factors such as rainfall explains the observed
pattern. East Africa Mountains are volcanic, young and bimodal rainfall pattern while the
Ethiopian Mountains were mostly forming by doming, are much older and have unimodal
pattern of rainfall.
The phenetic study, gross morphology and phylogenetic analysis based on two chloroplast
DNA (TrnL-F and rpL-16) and one nuclear DNA (ITS) reveal and confirm the existence of
two broadly distributed species viz: P. pictigluma and P. borussica and two narrow endemics,
namely P. chrysurus and P. dolichochaeta. Pentaschistis trisetoides is sister to the P.
pictigluma– P. borussica sub–clade. In addition the phylogenetic analysis of the Pentaschistis
clade provides a robust support for the two sub–clades of P. borussica–P. pictigluma and P.
chrysurus–P. dolichochaeta that indicate two events of independent colonization by these
two groups. Pentaschistis borussica can be easily distinguished from P. pictigluma, P. minor,
P. gracilis and P. mannii by its raised glands and open panicles. Both DNA sequence
similarity analyses as well as morphometric analysis among P. pictigluma, P. minor, P.
gracilis and P. minor indicate that these taxa are not distinct from each other and agree with
similar results obtained by S. M. Phillips. Hence this study proposes the retention of the
140
current taxonomic treatment that recognizes five species: P. borussica, P. pictigluma
(including the four varieties), P. trisetoides, P. chrysurus and P. dolichochaeta
.
4.2 Conclusions
The study supports the current taxonomic treatment that reduces P. minor, P. mannii, P.
gracilis to variety rank under P. pictigluma. The two narrow endemics similarly show close
morphological affinities, are older taxa undergoing range contraction and could be vulnerable
to natural and anthropogenic disturbances due to their limited distribution. They are also
distantly related to the two broadly distributed species, P. borussica and P. pictigluma.
Pentaschistis borussica and P. pictigluma species are young taxa that are undergoing rapid
diversification as is evident from their wide distribution range. Afroalpine vegetation types
are distinct and recognizable from each other but largely overlapping. Age, geographical
position, geological history, climate and edaphic factors account for the similarities and
disparities in species composition of the mountains and vegetation types studied.
4.3 Recommendations
This study recommends that;
•
Further population genetic studies be carried out among the populations of P.
pictigluma species complex and between the two narrow endemics as well as conduct
anatomical studies on leaf and glands to help further distinguish between the
populations of the broadly distributed and narrow endemic species.
•
Conservation programmes be initiated for the Afroalpine endemics since they are
vulnerable to anthropogenic disturbance and climate change.
•
Further research be carried out to determine why few families dominate the
Afroalpine vegetation communities.
141
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Appendix 1a: Data matrix of 12 plots of Mt Kenya showing presence/absence scores for all species sampled
Species
Adiantum thalictroides
Aeonium leucoblepharum
Agrocharis melanantha
Agrostis gracilifolia ssp. gracilifolia
Agrostis quinqueseta
Agrostis schimperiana
Agrostis sclerophylla
Agrostis trachyphylla
Agrostis volkensii
Aira caryophyllea
Alchemilla abyssinica
Alchemilla argyrophylla
Alchemilla haumannii
Alchemilla johnstonii
Alchemilla microbetula
Alchemilla pedata
Alchemilla subnivalis
Alchemilla stuhlmannii
Alchemilla triphylla
Alopecurus baptarrhenius
Anagallis serpens ssp. meyeri-johannis
Andropogon amethystinus
KER
P1
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
1
0
0
0
KER
P2
0
0
0
0
0
0
1
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
KER
P3
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
KER
P4
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
KEB
P1
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
KEB
P2
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
KEB
P3
0
0
0
1
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
KEB
P4
0
0
0
0
0
0
1
1
0
0
0
1
0
0
1
0
0
0
0
0
0
0
KED
P1
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
KED
P2
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
KED
P3
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
KED
P4
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
149
Andropogon lima
Anemone thomsonii var. thomsonii
Anogramma leptophylla
Anthemis tigrensis
Anthoxanthum nivale
Aphanes bachitii
Arabidopsis thaliana
Arabis alpine
Argyrolobium schimperianum
Artemisia afra
Asplenium abyssinicum
Asplenium aethiopicum
Asplenium buttneri
Barbarea intermedia
Bartsia decurva
Bartsia longiflora ssp. longiflora
Bromus leptoclados
Callitriche oreophila
Callitriche vulcanicola
Campanula edulis
Cardamine hirsuta
Cardamine obliqua
Carduus chamaecephalus
Carduus keniensis
Carduus leptacanthus
Carduus macracanthus
Carduus ruwenzoriensis
0
0
1
0
1
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
1
0
1
0
0
0
0
0
1
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
150
Carduus schimperi
Carex bequaerttii
Carex monostachya
Carex runssoroensis var. aberdarensis
Carex simensis
Carex conferta
Cerastium afromontanum
Cerastium octandrum
Cheilanthes farinosa
Cineraria abyssinica
Cineraria deltoidea
Colpodium chionogeiton
Colpodium hedbergii
Conyza spinosa
Conyza subscaposa
Cotula abyssinica
Cotula cryptocephala
Crassula granvikii
Crassula schimperi
Crassula sp.
Crepis dianthoseris
Crepis foetida
Crepis rueppellii
Cyanotis barbata
Cyanotis polyrrhiza
Cyperus elegantulus
Cyperus plateilema
1
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
1
0
0
1
0
0
1
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
1
0
0
0
0
0
0
1
0
1
0
0
0
0
1
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
1
0
0
0
0
1
0
0
0
0
1
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
151
Cystopteris fragilis
Delphinium macrocentrum
Delphinium wellbyi
Dendrosenecio adnivalis ssp. adnivalis
var. adnivalis
Dendrosenecio battiscombei
Dendrosenecio
elgonensis
ssp.
barbatipes
Dendrosenecio erici-rosenii ssp. ericirosenii
Dendrosenecio keniensis
Dendrosenecio
keniensis
×
keniodendron
Dendrosenecio keniodendron
Dendrosenecio kilimanjari ssp. cottonii
Deschampsia caespitosa
Deschampsia flexuosa var. afromontana
Dianthus longiglumis
Dichrocephala chrysanthemifolia
Dierama cupuliflorum
Dipsacus pinnatifidus
Disa stairsii
Echinops buhaitensis
Epilobium
stereophyllum
var.
stereophyllum
Erica arborea
Erica johnstonii
Erica tenuipilosa ssp. spicata
Erica trimera ssp. keniensis
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
1
0
1
0
1
0
1
0
0
0
1
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
1
0
0
1
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
152
Erigeron alpinus
Eriocaulon schimperi
Erophila verna
Euryops dacrydioides
Euryops elgonensis
Euryops prostratus
Festuca abyssinica
Festuca macrophylla
Festuca pilgeri
Festuca richardii
Festuca simensis
Galium acrophyum
Galium ruwenzoriense
Galium simense
Galium ossirwaense var. glabrum
Galium thunbergianum
Geranium arabicum
Geranium sp.nov.=Miehe 3002
Gnaphalium unionis
Haplocarpha rueppellii
Haplocarpha schimperi
Haplosciadium abyssinicum
Hebenstretia angolensis
Helichrysum brownei var. brownei
Helichrysum citrispinum
Helichrysum foetidum
Helichrysum formosissimum
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
1
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
1
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
153
Helichrysum forskahlii
Helichrysum globosum
Helichrysum gofense
Helichrysum horridum
Helichrysum kilimanjari
Helichrysum newii
Helichrysum ambylyphyllum
Helichrysum argyranthum
Helichrysum splendidum
Helichrysum stuhlmannii
Helictotrichon elongatum
Helictotrichon cf umbrosum
Heracleum abyssincum
Heracleum elgonense
Herniaria abyssinica
Hesperantha petitiana
Huperzia saururus
Hydrocotylee sibthorpioides
Hypericum peplidifolium
Hypericum revolutum
Hypericum afromontanum
Isolepis costata
Isolepis fluitans
Isolepis setacea
Kniphofia foliosa
Kniphofia isoetifolia
Kniphofia thomsonii
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
154
Koeleria capensis
Lactuca inermis
Limosella major
Lithospermum afromontanum
Lobelia aberdarica
Lobelia bequaertii
Lobelia deckenii
Lobelia gregoriana
Lobelia lindblomii
Lobelia rhynchopetalum
Lobelia stuhlmannii
Lobelia telekii
Lobelia wollastonii
Luzula abyssinica
Luzula johnstonii
Lychnis abyssinica
Lychnis rotundifolia
Malva verticillata
Merendera schimperiana
Minuartia filifolia
Myosotis keniensis
Myosotis vestergrenii
Oreophyton falcatum
Oxalis corniculata
Oxalis obliquifolia
Parietaria debilis
Paronychia bryoides
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
1
0
0
0
1
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
155
Pennisetum humile
Pentaschistis borussica
Pentaschistis pictigluma var gracilis
Pentaschistis pictigluma var. minor
Pentaschistis pictigluma var. pictigluma
Pentaschistis trisetoides
Peucedanum kerstenii
Phagnalon abyssinicum
Pimpinella oreophila
Pimpinella pimpinelloides
Plantago afra var. stricta
Poa annua
Poa leptoclada
Poa schimperiana
Polygala steudneri
Polypogon schimperianus
Polystichum magnificum
Polystichum setiferum
Ranunculus distrias
Ranunculus oligocarpus
Ranunculus oreophytus
Ranunculus stagnalis
Ranunculus trichophyllus
Ranunculus volkensii
Rhabdotosperma scrophulariifolia
Romulea fischeri
Rosularia semiensis
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
156
Rumex nepalensis
Rytidosperma grandiflorum
Rytidosperma subulata
Sagina abyssinica
Sagina afroalpina
Salvia nilotica
Satureja abyssinica
Satureja imbricata
Satureja kilimandschari
Satureja pseudosimensis
Satureja punctata ssp. punctata
Satureja simensis
Satureja uhligii var. obtusifolium
Saxifraga hederifolia
Scabiosa columbaria
Sedum crassularia
Sedum meyeri-johannis
Sedum mooneyi
Senecio balensis
Senecio farinaceus
Senecio fresenii
Senecio jacksonii
Senecio meyeri-johannis
Senecio nanus
Senecio polyadenus
Senecio purtschelleri
Senecio rhammatophyllus
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
157
Senecio schultzii
Senecio schweinfurthii
Senecio subsessilis
Senecio telekii
Senecio transmarinus
Senecio transmarinus var. major
Senecio transmarinus var. sycephyllus
Senecio unionis
Silene burchellii var. burchellii
Silene flammulifolia
Silene macrosolen
Silene melanolepis
Sonchus melanolepis
Stellaria media
Stoebe kilimandscharica
Subularia monticola
Swertia abyssinica
Swertia crassiuscula
Swertia engleri
Swertia kilimandscharica
Swertia pumila
Swertia subnivalis
Swertia uniflora
Swertia volkensii
Thymus schimperi
Thymus serrulatus
Trifolium acaule
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
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0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
158
Trifolium calocephalum
Trifolium cryptopodium
Trifolium elgonense
Trifolium multinerve
Trifolium rueppellianum
Trifolium simense
Umbilicus botryoides
Ursinia nana
Valerianella
microcarpa
microcarpa
Verbascum sedgwickianum
Veronica anagallis-aquatica
Veronica arvensis
Veronica glandulosa
Veronica gunae
Viola eminii
Vulpia bromoides
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
var.
159
Appendix 1b: Data matrix of 11 plots of Mt Elgon showing presence/absence scores for all species sampled
Species
Adiantum thalictroides
Aeonium leucoblepharum
Agrocharis melanantha
Agrostis gracilifolia ssp. gracilifolia
Agrostis quinqueseta
Agrostis schimperiana
Agrostis sclerophylla
Agrostis trachyphylla
Agrostis volkensii
Aira caryophyllea
Alchemilla abyssinica
Alchemilla argyrophylla
Alchemilla haumannii
Alchemilla johnstonii
Alchemilla microbetula
Alchemilla pedata
Alchemilla subnivalis
Alchemilla stuhlmannii
Alchemilla triphylla
Alopecurus baptarrhenius
Anagallis serpens ssp. meyeri-johannis
Andropogon amethystinus
ERP
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
ERP
2
0
0
0
1
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
ERP
3
0
0
0
1
0
0
0
0
0
0
0
1
0
1
1
0
0
0
0
0
0
0
ERP
4
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
EGP
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
EGP
2
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
EGP
3
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
EGP
4
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
1
0
EDP
1
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
EDP
2
0
0
0
1
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
EDP
3
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
160
Andropogon lima
Anemone thomsonii var. thomsonii
Anogramma leptophylla
Anthemis tigrensis
Anthoxanthum nivale
Aphanes bachitii
Arabidopsis thaliana
Arabis alpina
Argyrolobium schimperianum
Artemisia afra
Asplenium abyssinicum
Asplenium aethiopicum
Asplenium buttneri
Barbarea intermedia
Bartsia decurva
Bartsia longiflora ssp. longiflora
Bromus leptoclados
Callitriche oreophila
Callitriche vulcanicola
Campanula edulis
Cardamine hirsuta
Cardamine obliqua
Carduus chamaecephalus
Carduus keniensis
Carduus leptacanthus
Carduus macracanthus
Carduus ruwenzoriensis
0
0
0
0
1
0
0
1
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
1
0
0
0
1
0
0
0
0
0
1
0
1
0
0
0
1
0
0
0
0
0
0
1
0
1
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
161
Carduus schimperi
Carex bequaerttii
Carex monostachya
Carex runssoroensis var. aberdarensis
Carex simensis
Carex conferta
Cerastium afromontanum
Cerastium octandrum
Cheilanthes farinosa
Cineraria abyssinica
Cineraria deltoidea
Colpodium chionogeiton
Colpodium hedbergii
Conyza spinosa
Conyza subscaposa
Cotula abyssinica
Cotula cryptocephala
Crassula granvikii
Crassula schimperi
Crassula sp.
Crepis dianthoseris
Crepis foetida
Crepis rueppellii
Cyanotis barbata
Cyanotis polyrrhiza
Cyperus elegantulus
Cyperus plateilema
0
1
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
162
Cystopteris fragilis
0
Delphinium macrocentrum
0
Delphinium wellbyi
0
Dendrosenecio adnivalis ssp. adnivalis var.
0
adnivalis
Dendrosenecio battiscombei
0
Dendrosenecio elgonensis ssp. barbatipes
1
Dendrosenecio erici-rosenii ssp. erici-rosenii
0
Dendrosenecio keniensis
0
Dendrosenecio keniensis × keniodendron
0
Dendrosenecio keniodendron
0
Dendrosenecio kilimanjari ssp. cottonii
0
Deschampsia caespitosa
0
Deschampsia flexuosa var. afromontana
0
Dianthus longiglumis
0
Dichrocephala chrysanthemifolia
0
Dierama cupuliflorum
0
Dipsacus pinnatifidus
0
Disa stairsii
0
Echinops buhaitensis
0
Epilobium stereophyllum var. stereophyllum
0
Erica arborea
0
Erica johnstonii
0
Erica tenuipilosa ssp. spicata
0
Erica trimera ssp. keniensis
0
Erigeron alpinus
0
Eriocaulon schimperi
0
Erophila verna
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
163
Euryops dacrydioides
Euryops elgonensis
Euryops prostratus
Festuca abyssinica
Festuca macrophylla
Festuca pilgeri
Festuca richardii
Festuca simensis
Galium acrophyum
Galium ruwenzoriense
Galium simense
Galium ossirwaense var. glabrum
Galium thunbergianum
Geranium arabicum
Geranium sp.nov.=Miehe 3002
Gnaphalium unionis
Haplocarpha rueppellii
Haplocarpha schimperi
Haplosciadium abyssinicum
Hebenstretia angolensis
Helichrysum brownei var. brownei
Helichrysum citrispinum
Helichrysum foetidum
Helichrysum formosissimum
Helichrysum forskahlii
Helichrysum globosum
Helichrysum gofense
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
1
1
0
1
0
1
1
0
0
0
0
0
1
0
0
0
0
1
1
0
0
0
0
0
0
1
0
0
0
0
1
0
1
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
1
0
1
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
0
1
0
0
1
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
1
0
0
0
1
0
1
0
1
0
0
0
0
0
1
0
0
0
1
1
0
0
0
1
0
1
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
1
0
0
1
0
0
0
0
0
0
1
1
0
0
164
Helichrysum horridum
Helichrysum kilimanjari
Helichrysum newii
Helichrysum ambylyphyllum
Helichrysum argyranthum
Helichrysum splendidum
Helichrysum stuhlmannii
Helictotrichon elongatum
Helictotrichon cf umbrosum
Heracleum abyssincum
Heracleum elgonense
Herniaria abyssinica
Hesperantha petitiana
Huperzia saururus
Hydrocotylee sibthorpioides
Hypericum peplidifolium
Hypericum revolutum
Hypericum afromontanum
Isolepis costata
Isolepis fluitans
Isolepis setacea
Kniphofia foliosa
Kniphofia isoetifolia
Kniphofia thomsonii
Koeleria capensis
Lactuca inermis
Limosella major
0
1
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
1
1
1
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
165
Lithospermum afromontanum
Lobelia aberdarica
Lobelia bequaertii
Lobelia deckenii
Lobelia gregoriana
Lobelia lindblomii
Lobelia rhynchopetalum
Lobelia stuhlmannii
Lobelia telekii
Lobelia wollastonii
Luzula abyssinica
Luzula johnstonii
Lychnis abyssinica
Lychnis rotundifolia
Malva verticillata
Merendera schimperiana
Minuartia filifolia
Myosotis keniensis
Myosotis vestergrenii
Oreophyton falcatum
Oxalis corniculata
Oxalis obliquifolia
Parietaria debilis
Paronychia bryoides
Pennisetum humile
Pentaschistis borussica
Pentaschistis pictigluma var. gracilis
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
1
0
0
1
0
0
1
0
1
0
0
0
0
0
0
0
1
0
1
1
0
0
0
0
0
0
1
1
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
166
Pentaschistis pictigluma var. minor
Pentaschistis pictigluma var. pictigluma
Pentaschistis trisetoides
Peucedanum kerstenii
Phagnalon abyssinicum
Pimpinella oreophila
Pimpinella pimpinelloides
Plantago afra var. stricta
Poa annua
Poa leptoclada
Poa schimperiana
Polygala steudneri
Polypogon schimperianus
Polystichum magnificum
Polystichum setiferum
Ranunculus distrias
Ranunculus oligocarpus
Ranunculus oreophytus
Ranunculus stagnalis
Ranunculus trichophyllus
Ranunculus volkensii
Rhabdotosperma scrophulariifolia
Romulea fischeri
Rosularia semiensis
Rumex nepalensis
Rytidosperma grandiflorum
Rytidosperma subulata
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
167
Sagina abyssinica
Sagina afroalpina
Salvia nilotica
Satureja abyssinica
Satureja imbricata
Satureja kilimandschari
Satureja pseudosimensis
Satureja punctata ssp. punctata
Satureja simensis
Satureja uhligii var. obtusifolium
Saxifraga hederifolia
Scabiosa columbaria
Sedum crassularia
Sedum meyeri-johannis
Sedum mooneyi
Senecio balensis
Senecio farinaceus
Senecio fresenii
Senecio jacksonii
Senecio meyeri-johannis
Senecio nanus
Senecio polyadenus
Senecio purtschelleri
Senecio rhammatophyllus
Senecio schultzii
Senecio schweinfurthii
Senecio subsessilis
0
1
0
1
0
0
0
1
0
0
0
1
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
168
Senecio telekii
Senecio transmarinus
Senecio transmarinus var. major
Senecio transmarinus var. sycephyllus
Senecio unionis
Silene burchellii var. burchellii
Silene flammulifolia
Silene macrosolen
Silene melanolepis
Sonchus melanolepis
Stellaria media
Stoebe kilimandscharica
Subularia monticola
Swertia abyssinica
Swertia crassiuscula
Swertia engleri
Swertia kilimandscharica
Swertia pumila
Swertia subnivalis
Swertia uniflora
Swertia volkensii
Thymus schimperi
Thymus serrulatus
Trifolium acaule
Trifolium calocephalum
Trifolium cryptopodium
Trifolium elgonense
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
169
Trifolium multinerve
Trifolium rueppellianum
Trifolium simense
Umbilicus botryoides
Ursinia nana
Valerianella microcarpa var. microcarpa
Verbascum sedgwickianum
Veronica anagallis-aquatica
Veronica arvensis
Veronica glandulosa
Veronica gunae
Viola eminii
Vulpia bromoides
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
170
Appendix 1c :Data matrix of 12 plots of Mt Kilimanjaro showing presence/absence scores for all species sampled
Species
Adiantum thalictroides
Aeonium leucoblepharum
Agrocharis melanantha
Agrostis gracilifolia ssp. gracilifolia
Agrostis quinqueseta
Agrostis schimperiana
Agrostis sclerophylla
Agrostis trachyphylla
Agrostis volkensii
Aira caryophyllea
Alchemilla abyssinica
Alchemilla argyrophylla
Alchemilla haumannii
Alchemilla johnstonii
Alchemilla microbetula
Alchemilla pedata
Alchemilla subnivalis
Alchemilla stuhlmannii
Alchemilla triphylla
Alopecurus baptarrhenius
Anagallis serpens ssp. meyeri-johannis
Andropogon amethystinus
KIR
P1
0
0
0
0
0
0
0
0
1
0
0
1
0
1
0
0
0
0
0
0
0
0
KIR
P2
0
0
0
0
0
0
0
0
1
0
0
1
0
1
0
0
0
0
0
0
0
0
KIR
P3
0
0
0
0
0
0
0
0
1
0
0
1
0
1
0
0
0
0
0
0
1
0
KIR
P3
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
KIB
P1
0
0
0
0
0
0
0
0
1
0
0
1
0
0
1
0
0
0
0
0
1
0
KIB
P2
0
0
0
0
0
0
0
1
1
0
0
0
0
1
0
1
0
0
0
0
1
0
KIB
P3
0
0
0
0
0
0
0
1
0
0
0
1
0
1
0
0
0
0
0
0
0
0
KIB
P4
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
KIG
P1
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
1
0
KIG
P2
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
KIG
P3
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
KIG
P4
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
171
Andropogon lima
Anemone thomsonii var. thomsonii
Anogramma leptophylla
Anthemis tigrensis
Anthoxanthum nivale
Aphanes bachitii
Arabidopsis thaliana
Arabis alpina
Argyrolobium schimperianum
Artemisia afra
Asplenium abyssinicum
Asplenium aethiopicum
Asplenium buttneri
Barbarea intermedia
Bartsia decurva
Bartsia longiflora ssp. longiflora
Bromus leptoclados
Callitriche oreophila
Callitriche vulcanicola
Campanula edulis
Cardamine hirsuta
Cardamine obliqua
Carduus chamaecephalus
Carduus keniensis
Carduus leptacanthus
Carduus macracanthus
Carduus ruwenzoriensis
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
172
Carduus schimperi
Carex bequaerttii
Carex monostachya
Carex runssoroensis var. aberdarensis
Carex simensis
Carex conferta
Cerastium afromontanum
Cerastium octandrum
Cheilanthes farinosa
Cineraria abyssinica
Cineraria deltoidea
Colpodium chionogeiton
Colpodium hedbergii
Conyza spinosa
Conyza subscaposa
Cotula abyssinica
Cotula cryptocephala
Crassula granvikii
Crassula schimperi
Crassula sp.
Crepis dianthoseris
Crepis foetida
Crepis rueppellii
Cyanotis barbata
Cyanotis polyrrhiza
Cyperus elegantulus
Cyperus plateilema
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
173
Cystopteris fragilis
Delphinium macrocentrum
Delphinium wellbyi
Dendrosenecio adnivalis ssp. adnivalis
var. adnivalis
Dendrosenecio battiscombei
Dendrosenecio elgonensis ssp. barbatipes
Dendrosenecio erici-rosenii ssp. ericirosenii
Dendrosenecio keniensis
Dendrosenecio keniensis × keniodendron
Dendrosenecio keniodendron
Dendrosenecio kilimanjari ssp. cottonii
Deschampsia caespitosa
Deschampsia flexuosa var. afromontana
Dianthus longiglumis
Dichrocephala chrysanthemifolia
Dierama cupuliflorum
Dipsacus pinnatifidus
Disa stairsii
Echinops buhaitensis
Epilobium
stereophyllum
var.
stereophyllum
Erica arborea
Erica johnstonii
Erica tenuipilosa ssp. spicata
Erica trimera ssp. keniensis
Erigeron alpinus
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
1
0
0
1
0
0
1
0
0
0
0
0
1
0
0
1
0
0
1
1
0
1
0
0
1
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
1
1
0
0
0
0
1
1
0
0
0
0
0
1
0
0
1
174
Eriocaulon schimperi
Erophila verna
Euryops dacrydioides
Euryops elgonensis
Euryops prostratus
Festuca abyssinica
Festuca macrophylla
Festuca pilgeri
Festuca richardii
Festuca simensis
Galium acrophyum
Galium ruwenzoriense
Galium simense
Galium ossirwaense var. glabrum
Galium thunbergianum
Geranium arabicum
Geranium sp.nov.=Miehe 3002
Gnaphalium unionis
Haplocarpha rueppellii
Haplocarpha schimperi
Haplosciadium abyssinicum
Hebenstretia angolensis
Helichrysum brownei var. brownei
Helichrysum citrispinum
Helichrysum foetidum
Helichrysum formosissimum
Helichrysum forskahlii
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
1
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
1
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
1
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
1
0
0
0
0
1
0
0
0
1
0
1
0
0
0
0
1
0
0
0
0
0
0
0
1
0
1
0
0
1
0
0
1
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
1
0
0
1
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
1
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
175
Helichrysum globosum
Helichrysum gofense
Helichrysum horridum
Helichrysum kilimanjari
Helichrysum newii
Helichrysum ambylyphyllum
Helichrysum argyranthum
Helichrysum splendidum
Helichrysum stuhlmannii
Helictotrichon elongatum
Helictotrichon cf umbrosum
Heracleum abyssincum
Heracleum elgonense
Herniaria abyssinica
Hesperantha petitiana
Huperzia saururus
Hydrocotylee sibthorpioides
Hypericum peplidifolium
Hypericum revolutum
Hypericum afromontanum
Isolepis costata
Isolepis fluitans
Isolepis setacea
Kniphofia foliosa
Kniphofia isoetifolia
Kniphofia thomsonii
Koeleria capensis
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
1
176
Lactuca inermis
Limosella major
Lithospermum afromontanum
Lobelia aberdarica
Lobelia bequaertii
Lobelia deckenii
Lobelia gregoriana
Lobelia lindblomii
Lobelia rhynchopetalum
Lobelia stuhlmannii
Lobelia telekii
Lobelia wollastonii
Luzula abyssinica
Luzula johnstonii
Lychnis abyssinica
Lychnis rotundifolia
Malva verticillata
Merendera schimperiana
Minuartia filifolia
Myosotis keniensis
Myosotis vestergrenii
Oreophyton falcatum
Oxalis corniculata
Oxalis obliquifolia
Parietaria debilis
Paronychia bryoides
Pennisetum humile
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
1
0
1
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
177
Pentaschistis borussica
Pentaschistis pictigluma var gracilis
Pentaschistis pictigluma var. minor
Pentaschistis pictigluma var. pictigluma
Pentaschistis trisetoides
Peucedanum kerstenii
Phagnalon abyssinicum
Pimpinella oreophila
Pimpinella pimpinelloides
Plantago afra var. stricta
Poa annua
Poa leptoclada
Poa schimperiana
Polygala steudneri
Polypogon schimperianus
Polystichum magnificum
Polystichum setiferum
Ranunculus distrias
Ranunculus oligocarpus
Ranunculus oreophytus
Ranunculus stagnalis
Ranunculus trichophyllus
Ranunculus volkensii
Rhabdotosperma scrophulariifolia
Romulea fischeri
Rosularia semiensis
Rumex nepalensis
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
178
Rytidosperma grandiflorum
Rytidosperma subulata
Sagina abyssinica
Sagina afroalpina
Salvia nilotica
Satureja abyssinica
Satureja imbricata
Satureja kilimandschari
Satureja pseudosimensis
Satureja punctata ssp. punctata
Satureja simensis
Satureja uhligii var. obtusifolium
Saxifraga hederifolia
Scabiosa columbaria
Sedum crassularia
Sedum meyeri-johannis
Sedum mooneyi
Senecio balensis
Senecio farinaceus
Senecio fresenii
Senecio jacksonii
Senecio meyeri-johannis
Senecio nanus
Senecio polyadenus
Senecio purtschelleri
Senecio rhammatophyllus
Senecio schultzii
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
179
Senecio schweinfurthii
Senecio subsessilis
Senecio telekii
Senecio transmarinus
Senecio transmarinus var. major
Senecio transmarinus var. sycephyllus
Senecio unionis
Silene burchellii var. burchellii
Silene flammulifolia
Silene macrosolen
Silene melanolepis
Sonchus melanolepis
Stellaria media
Stoebe kilimandscharica
Subularia monticola
Swertia abyssinica
Swertia crassiuscula
Swertia engleri
Swertia kilimandscharica
Swertia pumila
Swertia subnivalis
Swertia uniflora
Swertia volkensii
Thymus schimperi
Thymus serrulatus
Trifolium acaule
Trifolium calocephalum
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
180
Trifolium cryptopodium
Trifolium elgonense
Trifolium multinerve
Trifolium rueppellianum
Trifolium simense
Umbilicus botryoides
Ursinia nana
Valerianella microcarpa var. microcarpa
Verbascum sedgwickianum
Veronica anagallis-aquatica
Veronica arvensis
Veronica glandulosa
Veronica gunae
Viola eminii
Vulpia bromoides
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
181
Appendix 1d: Data matrix of 12 plots of Mt Rwenzori showing presence/absence scores for all species sampled
Species
Adiantum thalictroides
Aeonium leucoblepharum
Agrocharis melanantha
Agrostis gracilifolia ssp. gracilifolia
Agrostis quinqueseta
Agrostis schimperiana
Agrostis sclerophylla
Agrostis trachyphylla
Agrostis volkensii
Aira caryophyllea
Alchemilla abyssinica
Alchemilla argyrophylla
Alchemilla haumannii
Alchemilla johnstonii
Alchemilla microbetula
Alchemilla pedata
Alchemilla subnivalis
Alchemilla stuhlmannii
Alchemilla triphylla
Alopecurus baptarrhenius
Anagallis serpens ssp. meyeri-johannis
Andropogon amethystinus
Andropogon lima
RBP
1
0
0
1
0
0
1
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
RBP
2
0
0
0
0
0
1
0
0
0
0
0
1
0
1
1
0
0
0
0
0
0
0
0
RBP
3
0
0
0
0
0
0
0
0
0
0
0
1
0
1
1
0
0
0
0
0
0
0
0
RBP
3
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
RDP
1
0
0
0
0
0
0
0
0
0
0
0
1
0
1
1
0
0
1
1
0
0
0
0
RDP
2
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
RDP
3
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
RDP
4
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
1
1
0
0
0
0
RAP
1
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
RAP
2
0
0
0
0
0
0
0
0
0
0
0
1
0
1
1
0
0
0
1
0
0
0
0
RAP
3
0
0
0
0
0
0
0
0
0
0
0
1
0
1
1
0
0
0
1
0
0
0
0
RAP
4
0
0
0
0
0
1
0
0
0
0
0
1
0
1
1
0
0
1
1
0
0
0
0
182
Anemone thomsonii var. thomsonii
Anogramma leptophylla
Anthemis tigrensis
Anthoxanthum nivale
Aphanes bachitii
Arabidopsis thaliana
Arabis alpina
Argyrolobium schimperianum
Artemisia afra
Asplenium abyssinicum
Asplenium aethiopicum
Asplenium buttneri
Barbarea intermedia
Bartsia decurva
Bartsia longiflora ssp. longiflora
Bromus leptoclados
Callitriche oreophila
Callitriche vulcanicola
Campanula edulis
Cardamine hirsuta
Cardamine obliqua
Carduus chamaecephalus
Carduus keniensis
Carduus leptacanthus
Carduus macracanthus
Carduus ruwenzoriensis
Carduus schimperi
0
0
0
1
0
0
1
0
0
0
1
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
183
Carex bequaerttii
Carex monostachya
Carex runssoroensis var. aberdarensis
Carex simensis
Carex conferta
Cerastium afromontanum
Cerastium octandrum
Cheilanthes farinosa
Cineraria abyssinica
Cineraria deltoidea
Colpodium chionogeiton
Colpodium hedbergii
Conyza spinosa
Conyza subscaposa
Cotula abyssinica
Cotula cryptocephala
Crassula granvikii
Crassula schimperi
Crassula sp.
Crepis dianthoseris
Crepis foetida
Crepis rueppellii
Cyanotis barbata
Cyanotis polyrrhiza
Cyperus elegantulus
Cyperus plateilema
Cystopteris fragilis
1
0
1
0
0
0
1
1
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
184
Delphinium macrocentrum
Delphinium wellbyi
Dendrosenecio adnivalis ssp. adnivalis var.
adnivalis
Dendrosenecio battiscombei
Dendrosenecio elgonensis ssp. barbatipes
Dendrosenecio erici-rosenii ssp. ericirosenii
Dendrosenecio keniensis
Dendrosenecio keniensis × keniodendron
Dendrosenecio keniodendron
Dendrosenecio kilimanjari ssp. cottonii
Deschampsia caespitosa
Deschampsia flexuosa var. afromontana
Dianthus longiglumis
Dichrocephala chrysanthemifolia
Dierama cupuliflorum
Dipsacus pinnatifidus
Disa stairsii
Echinops buhaitensis
Epilobium
stereophyllum
var.
stereophyllum
Erica arborea
Erica johnstonii
Erica tenuipilosa ssp. spicata
Erica trimera ssp. keniensis
Erigeron alpinus
Eriocaulon schimperi
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
1
0
0
1
0
0
1
0
0
1
0
0
1
0
0
1
0
0
1
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
1
1
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
185
Erophila verna
Euryops dacrydioides
Euryops elgonensis
Euryops prostratus
Festuca abyssinica
Festuca macrophylla
Festuca pilgeri
Festuca richardii
Festuca simensis
Galium acrophyum
Galium ruwenzoriense
Galium simense
Galium ossirwaense var. glabrum
Galium thunbergianum
Geranium arabicum
Geranium sp.nov.=Miehe 3002
Gnaphalium unionis
Haplocarpha rueppellii
Haplocarpha schimperi
Haplosciadium abyssinicum
Hebenstretia angolensis
Helichrysum brownei var. brownei
Helichrysum citrispinum
Helichrysum foetidum
Helichrysum formosissimum
Helichrysum forskahlii
Helichrysum globosum
0
0
0
0
1
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
186
Helichrysum gofense
Helichrysum horridum
Helichrysum kilimanjari
Helichrysum newii
Helichrysum ambylyphyllum
Helichrysum argyranthum
Helichrysum splendidum
Helichrysum stuhlmannii
Helictotrichon elongatum
Helictotrichon cf umbrosum
Heracleum abyssincum
Heracleum elgonense
Herniaria abyssinica
Hesperantha petitiana
Huperzia saururus
Hydrocotylee sibthorpioides
Hypericum peplidifolium
Hypericum revolutum
Hypericum afromontanum
Isolepis costata
Isolepis fluitans
Isolepis setacea
Kniphofia foliosa
Kniphofia isoetifolia
Kniphofia thomsonii
Koeleria capensis
Lactuca inermis
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
1
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
187
Limosella major
Lithospermum afromontanum
Lobelia aberdarica
Lobelia bequaertii
Lobelia deckenii ssp. deckenii, ssp. burttii
Lobelia gregoriana ssp. elgonensis
Lobelia lindblomii
Lobelia rhynchopetalum
Lobelia stuhlmannii
Lobelia telekii
Lobelia wollastonii
Luzula abyssinica
Luzula johnstonii
Lychnis abyssinica
Lychnis rotundifolia
Malva verticillata
Merendera schimperiana
Minuartia filifolia
Myosotis keniensis
Myosotis vestergrenii
Oreophyton falcatum
Oxalis corniculata
Oxalis obliquifolia
Parietaria debilis
Paronychia bryoides
Pennisetum humile
Pentaschistis borussica
1
0
0
1
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
188
Pentaschistis pictigluma var. gracilis
Pentaschistis pictigluma var. minor
Pentaschistis pictigluma var. pictigluma
Pentaschistis trisetoides
Peucedanum kerstenii
Phagnalon abyssinicum
Pimpinella oreophila
Pimpinella pimpinelloides
Plantago afra var. stricta
Poa annua
Poa leptoclada
Poa schimperiana
Polygala steudneri
Polypogon schimperianus
Polystichum magnificum
Polystichum setiferum
Ranunculus distrias
Ranunculus oligocarpus
Ranunculus oreophytus
Ranunculus stagnalis
Ranunculus trichophyllus
Ranunculus volkensii
Rhabdotosperma scrophulariifolia
Romulea fischeri
Rosularia semiensis
Rumex nepalensis
Rytidosperma grandiflorum
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
1
1
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
189
Rytidosperma subulata
Sagina abyssinica
Sagina afroalpina
Salvia nilotica
Satureja abyssinica
Satureja imbricata
Satureja kilimandschari
Satureja pseudosimensis
Satureja punctata ssp. punctata
Satureja simensis
Satureja uhligii var. obtusifolium
Saxifraga hederifolia
Scabiosa columbaria
Sedum crassularia
Sedum meyeri-johannis
Sedum mooneyi
Senecio balensis
Senecio farinaceus
Senecio fresenii
Senecio jacksonii
Senecio meyeri-johannis
Senecio nanus
Senecio polyadenus
Senecio purtschelleri
Senecio rhammatophyllus
Senecio schultzii
Senecio schweinfurthii
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
190
Senecio subsessilis
Senecio telekii
Senecio transmarinus
Senecio transmarinus var. major
Senecio transmarinus var. sycephyllus
Senecio unionis
Silene burchellii var burchellii
Silene flammulifolia
Silene macrosolen
Silene melanolepis
Sonchus melanolepis
Stellaria media
Stoebe kilimandscharica
Subularia monticola
Swertia abyssinica
Swertia crassiuscula
Swertia engleri var. engleri
Swertia kilimandscharica
Swertia pumila
Swertia subnivalis
Swertia uniflora
Swertia volkensii
Thymus schimperi
Thymus serrulatus
Trifolium acaule
Trifolium calocephalum
Trifolium cryptopodium
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
191
Trifolium elgonense
Trifolium multinerve
Trifolium rueppellianum
Trifolium simense
Umbilicus botryoides
Ursinia nana
Valerianella microcarpa,var. microcarpa
Verbascum sedgwickianum
Veronica anagallis-aquatica
Veronica arvensis
Veronica glandulosa
Veronica gunae
Viola eminii
Vulpia bromoides
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
192
Appendix 1e: Data matrix of 16 plots of Mt Bale showing presence/absence scores for all species sampled
Species
Adiantum thalictroides
Aeonium leucoblepharum
Agrocharis melanantha
Agrostis gracilifolia ssp.
gracilifolia
Agrostis quinqueseta
Agrostis schimperiana
Agrostis sclerophylla
Agrostis trachyphylla
Agrostis volkensii
Aira caryophyllea
Alchemilla abyssinica
Alchemilla argyrophylla
Alchemilla haumannii
Alchemilla johnstonii
Alchemilla microbetula
Alchemilla pedata
Alchemilla subnivalis
Alchemilla stuhlmannii
Alchemilla triphylla
Alopecurus baptarrhenius
Anagallis
serpens
ssp.
meyeri-johannis
Andropogon amethystinus
Andropogon lima
BRP
1
BRP
2
BRP
3
BRP
4
BBP
1
BBP
2
BBP
3
BBP
4
BGP
1
BGP
2
BGP
3
BGP
4
BAP
1
BAP
2
BAP
3
BAP
4
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
1
1
0
0
1
1
1
1
0
1
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
1
0
1
1
0
1
0
0
1
1
0
0
0
0
0
0
0
1
1
0
1
0
1
1
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
1
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
193
Anemone thomsonii var.
thomsonii
Anogramma leptophylla
Anthemis tigrensis
Anthoxanthum nivale
Aphanes bachitii
Arabidopsis thaliana
Arabis alpina
Argyrolobium
schimperianum
Artemisia afra
Asplenium abyssinicum
Asplenium aethiopicum
Asplenium buttneri
Barbarea intermedia
Bartsia decurva
Bartsia
longiflora
ssp.
longiflora
Bromus leptoclados
Callitriche oreophila
Callitriche vulcanicola
Campanula edulis
Cardamine hirsuta
Cardamine oblique
Carduus chamaecephalus
Carduus keniensis
Carduus leptacanthus
Carduus macracanthus
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
1
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
1
1
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
194
Carduus ruwenzoriensis
Carduus schimperi
Carex bequaerttii
Carex monostachya
Carex runssoroensis var.
aberdarensis
Carex simensis
Carex conferta
Cerastium afromontanum
Cerastium octandrum
Cheilanthes farinosa
Cineraria abyssinica
Cineraria deltoidea
Colpodium chionogeiton
Colpodium hedbergii
Conyza spinosa
Conyza subscaposa
Cotula abyssinica
Cotula cryptocephala
Crassula granvikii
Crassula schimperi
Crassula sp.
Crepis dianthoseris
Crepis foetida
Crepis rueppellii
Cyanotis barbata
Cyanotis polyrrhiza
Cyperus elegantulus
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
0
0
1
0
0
1
1
1
1
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
1
1
1
1
1
1
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
1
0
0
0
1
0
1
1
1
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
0
1
1
1
1
0
1
0
1
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
195
Cyperus plateilema
Cystopteris fragilis
Delphinium macrocentrum
Delphinium wellbyi
Dendrosenecio
adnivalis
ssp. adnivalis var. adnivalis
Dendrosenecio battiscombei
Dendrosenecio elgonensis
ssp. barbatipes
Dendrosenecio erici-rosenii
ssp. erici-rosenii
Dendrosenecio keniensis
Dendrosenecio keniensis ×
keniodendron
Dendrosenecio
keniodendron
Dendrosenecio kilimanjari
ssp. cottonii
Deschampsia caespitosa
Deschampsia flexuosa var.
afromontana
Dianthus longiglumis
Dichrocephala
chrysanthemifolia
Dierama cupuliflorum
Dipsacus pinnatifidus
Disa stairsii
Echinops buhaitensis
Epilobium
stereophyllum
var. stereophyllum
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
1
0
1
0
1
1
1
1
1
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
196
Erica arborea
Erica johnstonii
Erica
tenuipilosa
ssp.
spicata
Erica trimera ssp. keniensis
Erigeron alpines
Eriocaulon schimperi
Erophila verna
Euryops dacrydioides
Euryops elgonensis
Euryops prostrates
Festuca abyssinica
Festuca macrophylla
Festuca pilgeri
Festuca richardii
Festuca simensis
Galium acrophyum
Galium ruwenzoriense
Galium simense
Galium ossirwaense var.
glabrum
Galium thunbergianum
Geranium arabicum
Geranium
sp.nov.=Miehe
3002
Gnaphalium unionis
Haplocarpha rueppellii
Haplocarpha schimperi
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
1
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
1
1
1
1
1
0
1
1
1
0
0
0
0
0
0
1
1
1
1
0
1
1
1
1
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
1
1
0
0
0
0
1
0
0
0
0
0
0
1
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
1
1
1
1
1
0
0
0
0
1
1
1
1
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
197
Haplosciadium abyssinicum
Hebenstretia angolensis
Helichrysum brownei var.
brownei
Helichrysum citrispinum
Helichrysum foetidum
Helichrysum formosissimum
Helichrysum forskahlii
Helichrysum globosum
Helichrysum gofense
Helichrysum horridum
Helichrysum kilimanjari
Helichrysum newii
Helichrysum ambylyphyllum
Helichrysum argyranthum
Helichrysum splendidum
Helichrysum stuhlmannii
Helictotrichon elongatum
Helictotrichon cf umbrosum
Heracleum abyssincum
Heracleum elgonense
Herniaria abyssinica
Hesperantha petitiana
Huperzia saururus
Hydrocotylee sibthorpioides
Hypericum peplidifolium
Hypericum revolutum
Hypericum afromontanum
0
0
1
0
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
1
0
1
0
0
1
1
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
1
1
0
1
0
1
0
0
1
1
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
1
1
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
1
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
198
Isolepis costata
Isolepis fluitans
Isolepis setacea
Kniphofia foliosa
Kniphofia isoetifolia
Kniphofia thomsonii
Koeleria capensis
Lactuca inermis
Limosella major
Lithospermum
afromontanum
Lobelia aberdarica
Lobelia bequaertii
Lobelia
deckenii
ssp.
deckenii, ssp. burttii
Lobelia gregoriana ssp.
elgonensis, gregoriana
Lobelia lindblomii
Lobelia rhynchopetalum
Lobelia stuhlmannii
Lobelia telekii
Lobelia wollastonii
Luzula abyssinica
Luzula johnstonii
Lychnis abyssinica
Lychnis rotundifolia
Malva verticillata
Merendera schimperiana
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
0
0
0
0
0
0
1
1
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
1
1
1
1
1
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
1
0
1
0
1
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
1
1
1
1
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
199
Minuartia filifolia
Myosotis keniensis
Myosotis vestergrenii
Oreophyton falcatum
Oxalis corniculata
Oxalis obliquifolia
Parietaria debilis
Paronychia bryoides
Pennisetum humile
Pentaschistis borussica
Pentaschistis pictigluma var.
gracilis
Pentaschistis pictigluma var.
minor
Pentaschistis pictigluma var.
pictigluma
Pentaschistis trisetoides
Peucedanum kerstenii
Phagnalon abyssinicum
Pimpinella oreophila
Pimpinella pimpinelloides
Plantago afra var. stricta
Poa annua
Poa leptoclada
Poa schimperiana
Polygala steudneri
Polypogon schimperianus
Polystichum magnificum
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
1
1
0
0
1
1
0
1
0
1
0
0
0
0
1
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
0
1
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
1
0
1
1
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
200
Polystichum setiferum
Ranunculus distrias
Ranunculus oligocarpus
Ranunculus oreophytus
Ranunculus stagnalis
Ranunculus trichophyllus
Ranunculus volkensii
Rhabdotosperma
scrophulariifolia
Romulea fischeri
Rosularia semiensis
Rumex nepalensis
Rytidosperma grandiflorum
Rytidosperma subulata
Sagina abyssinica
Sagina afroalpina
Salvia nilotica
Satureja abyssinica
Satureja imbricate
Satureja kilimandschari
Satureja pseudosimensis
Satureja
punctata
ssp.
punctata
Satureja simensis
Satureja
uhligii
var.
obtusifolium
Saxifraga hederifolia
Scabiosa columbaria
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
0
0
0
1
1
1
1
0
0
1
1
0
0
0
0
0
1
0
0
0
0
1
0
1
1
0
0
1
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
201
Sedum crassularia
Sedum meyeri-johannis
Sedum mooneyi
Senecio balensis
Senecio farinaceous
Senecio fresenii
Senecio jacksonii
Senecio meyeri-johannis
Senecio nanus
Senecio polyadenus
Senecio purtschelleri
Senecio rhammatophyllus
Senecio schultzii
Senecio schweinfurthii
Senecio subsessilis
Senecio telekii
Senecio transmarinus
Senecio transmarinus var.
major
Senecio transmarinus var.
sycephyllus
Senecio unionis
Silene
burchellii
var.
burchellii
Silene flammulifolia
Silene macrosolen
Silene melanolepis
Sonchus melanolepis
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
202
Stellaria media
Stoebe kilimandscharica
Subularia monticola
Swertia abyssinica
Swertia crassiuscula
Swertia engleri var. engleri
Swertia kilimandscharica
Swertia pumila
Swertia subnivalis
Swertia uniflora
Swertia volkensii
Thymus schimperi
Thymus serrulatus
Trifolium acaule
Trifolium calocephalum
Trifolium cryptopodium
Trifolium elgonense
Trifolium multinerve
Trifolium rueppellianum
Trifolium simense
Umbilicus botryoides
Ursinia nana
Valerianella
microcarpa
var. microcarpa
Verbascum sedgwickianum
Veronica anagallis-aquatica
Veronica arvensis
Veronica glandulosa
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
1
0
1
1
1
1
1
1
0
0
0
0
0
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
1
0
0
1
1
1
0
0
0
1
1
0
0
203
Veronica gunae
Viola eminii
Vulpia bromoides
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
204
Appendix 1f: Data matrix of 12 plots of Mt Simen showing presence/absence scores for all species sampled
Species
Adiantum thalictroides
Aeonium leucoblepharum
Agrocharis melanantha
Agrostis gracilifolia ssp. gracilifolia
Agrostis quinqueseta
Agrostis schimperiana
Agrostis sclerophylla
Agrostis trachyphylla
Agrostis volkensii
Aira caryophyllea
Alchemilla abyssinica
Alchemilla argyrophylla
Alchemilla haumannii
Alchemilla johnstonii
Alchemilla microbetula
Alchemilla pedata
Alchemilla subnivalis
Alchemilla stuhlmannii
Alchemilla triphylla
Alopecurus baptarrhenius
Anagallis serpens ssp. meyerijohannis
Andropogon amethystinus
Andropogon lima
SRP1
SRP2
SRP3
SRP3
SBP1
SBP2
SBP3
SBP4
SGP1
SGP2
SGP3
SGP4
1
1
1
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
1
0
0
1
0
1
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
1
0
0
1
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
1
0
0
1
0
0
0
1
0
0
1
0
0
1
0
0
1
0
0
0
1
1
0
0
0
0
0
0
1
0
0
205
Anemone thomsonii var. thomsonii
Anogramma leptophylla
Anthemis tigrensis
Anthoxanthum nivale
Aphanes bachitii
Arabidopsis thaliana
Arabis alpine
Argyrolobium schimperianum
Artemisia afra
Asplenium abyssinicum
Asplenium aethiopicum
Asplenium buttneri
Barbarea intermedia
Bartsia decurva
Bartsia longiflora ssp. longiflora
Bromus leptoclados
Callitriche oreophila
Callitriche vulcanicola
Campanula edulis
Cardamine hirsuta
Cardamine obliqua
Carduus chamaecephalus
Carduus keniensis
Carduus leptacanthus
Carduus macracanthus
Carduus ruwenzoriensis
Carduus schimperi
0
1
0
0
0
1
1
0
0
0
1
0
0
0
1
0
0
0
1
1
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
0
1
1
0
0
0
0
1
0
0
1
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
1
0
0
1
0
1
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
0
0
1
0
0
1
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
1
0
0
0
1
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
206
Carex bequaerttii
Carex monostachya
Carex
runssoroensis
aberdarensis
Carex simensis
Carex conferta
Cerastium afromontanum
Cerastium octandrum
Cheilanthes farinosa
Cineraria abyssinica
Cineraria deltoidea
Colpodium chionogeiton
Colpodium hedbergii
Conyza spinosa
Conyza subscaposa
Cotula abyssinica
Cotula cryptocephala
Crassula granvikii
Crassula schimperi
Crassula sp.
Crepis dianthoseris
Crepis foetida
Crepis rueppellii
Cyanotis barbata
Cyanotis polyrrhiza
Cyperus elegantulus
Cyperus plateilema
Cystopteris fragilis
0
0
0
0
0
0
0
0
0
1
0
1
0
1
0
1
0
0
0
0
0
1
0
1
0
0
0
0
1
0
0
1
0
1
1
0
0
0
1
1
0
1
0
1
0
1
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
1
0
1
1
0
1
1
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
1
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
1
0
1
1
0
1
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
1
0
1
0
0
1
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
1
0
0
0
1
1
0
0
1
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
1
0
0
1
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
1
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
1
0
0
1
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
1
1
0
1
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
var.
207
Delphinium macrocentrum
Delphinium wellbyi
Dendrosenecio
adnivalis
ssp.
adnivalis var. adnivalis
Dendrosenecio battiscombei
Dendrosenecio elgonensis ssp.
barbatipes
Dendrosenecio erici-rosenii ssp.
erici-rosenii
Dendrosenecio keniensis
Dendrosenecio
keniensis
×
keniodendron
Dendrosenecio keniodendron
Dendrosenecio kilimanjari ssp.
cottonii
Deschampsia caespitosa
Deschampsia
flexuosa
var.
afromontana
Dianthus longiglumis
Dichrocephala chrysanthemifolia
Dierama cupuliflorum
Dipsacus pinnatifidus
Disa stairsii
Echinops buhaitensis
Epilobium
stereophyllum
var.
stereophyllum
Erica arborea
Erica johnstonii
Erica tenuipilosa ssp. spicata
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
1
0
0
0
1
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
1
0
0
0
1
0
0
0
1
0
0
0
0
1
0
0
0
0
0
1
0
1
0
1
0
0
0
1
208
Erica trimera ssp. keniensis
Erigeron alpines
Eriocaulon schimperi
Erophila verna
Euryops dacrydioides
Euryops elgonensis
Euryops prostrates
Festuca abyssinica
Festuca macrophylla
Festuca pilgeri
Festuca richardii
Festuca simensis
Galium acrophyum
Galium ruwenzoriense
Galium simense
Galium ossirwaense var. glabrum
Galium thunbergianum
Geranium arabicum
Geranium sp.nov.=Miehe 3002
Gnaphalium unionis
Haplocarpha rueppellii
Haplocarpha schimperi
Haplosciadium abyssinicum
Hebenstretia angolensis
Helichrysum brownei var. brownei
Helichrysum citrispinum
Helichrysum foetidum
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
1
0
0
1
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
1
1
0
1
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
1
1
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
1
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
209
Helichrysum formosissimum
Helichrysum forskahlii
Helichrysum globosum
Helichrysum gofense
Helichrysum horridum
Helichrysum kilimanjari
Helichrysum newii
Helichrysum ambylyphyllum
Helichrysum argyranthum
Helichrysum splendidum
Helichrysum stuhlmannii
Helictotrichon elongatum
Helictotrichon cf umbrosum
Heracleum abyssincum
Heracleum elgonense
Herniaria abyssinica
Hesperantha petitiana
Huperzia saururus
Hydrocotylee sibthorpioides
Hypericum peplidifolium
Hypericum revolutum
Hypericum afromontanum
Isolepis costata
Isolepis fluitans
Isolepis setacea
Kniphofia foliosa
Kniphofia isoetifolia
0
0
0
0
1
0
0
0
0
0
0
1
0
1
0
1
1
0
0
0
0
0
0
0
1
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
1
0
0
0
0
0
1
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
210
Kniphofia thomsonii
Koeleria capensis
Lactuca inermis
Limosella major
Lithospermum afromontanum
Lobelia aberdarica
Lobelia bequaertii
Lobelia deckenii ssp. deckenii/ ssp.
burttii
Lobelia gregoriana ssp.elgonensis/
ssp.gregoriana
Lobelia lindblomii
Lobelia rhynchopetalum
Lobelia stuhlmannii
Lobelia telekii
Lobelia wollastonii
Luzula abyssinica
Luzula johnstonii
Lychnis abyssinica
Lychnis rotundifolia
Malva verticillata
Merendera schimperiana
Minuartia filifolia
Myosotis keniensis
Myosotis vestergrenii
Oreophyton falcatum
Oxalis corniculata
Oxalis obliquifolia
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
1
0
1
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
1
1
0
0
0
0
1
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
211
Parietaria debilis
Paronychia bryoides
Pennisetum humile
Pentaschistis borussica
Pentaschistis
pictigluma
var.
gracilis
Pentaschistis pictigluma var. minor
Pentaschistis
pictigluma
var.
pictigluma
Pentaschistis trisetoides
Peucedanum kerstenii
Phagnalon abyssinicum
Pimpinella oreophila
Pimpinella pimpinelloides
Plantago afra var. stricta
Poa annua
Poa leptoclada
Poa schimperiana
Polygala steudneri
Polypogon schimperianus
Polystichum magnificum
Polystichum setiferum
Ranunculus distrias
Ranunculus oligocarpus
Ranunculus oreophytus
Ranunculus stagnalis
Ranunculus trichophyllus
Ranunculus volkensii
0
0
0
0
1
0
0
0
1
0
0
0
1
1
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
1
0
0
1
1
1
1
0
0
1
0
0
0
1
0
0
0
0
0
0
1
0
0
0
1
1
1
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
1
1
1
1
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
1
1
1
0
0
0
0
0
0
0
0
1
0
0
0
1
1
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
212
Rhabdotosperma scrophulariifolia
Romulea fischeri
Rosularia semiensis
Rumex nepalensis
Rytidosperma grandiflorum
Rytidosperma subulata
Sagina abyssinica
Sagina afroalpina
Salvia nilotica
Satureja abyssinica
Satureja imbricata
Satureja kilimandschari
Satureja pseudosimensis
Satureja punctata ssp. punctata
Satureja simensis
Satureja uhligii var. obtusifolium
Saxifraga hederifolia
Scabiosa columbaria
Sedum crassularia
Sedum meyeri-johannis
Sedum mooneyi
Senecio balensis
Senecio farinaceous
Senecio fresenii
Senecio jacksonii
Senecio meyeri-johannis
Senecio nanus
1
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
1
0
1
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
1
0
1
0
0
1
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
1
0
0
1
0
1
1
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
1
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
1
0
1
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
213
Senecio polyadenus
Senecio purtschelleri
Senecio rhammatophyllus
Senecio schultzii
Senecio schweinfurthii
Senecio subsessilis
Senecio telekii
Senecio transmarinus
Senecio transmarinus var. major
Senecio
transmarinus
var.
sycephyllus
Senecio unionis
Silene burchellii var. burchellii
Silene flammulifolia
Silene macrosolen
Silene melanolepis
Sonchus melanolepis
Stellaria media
Stoebe kilimandscharica
Subularia monticola
Swertia abyssinica
Swertia
crassiuscula
ssp.crassiuscula, ssp. robusta
Swertia engleri var. engleri (ssp.
engleri,ssp. woodii)
Swertia kilimandscharica
Swertia pumila
Swertia subnivalis
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
1
0
0
0
0
0
0
1
0
0
0
1
1
0
0
0
1
0
0
0
0
1
1
1
0
0
0
0
0
0
1
1
0
0
1
1
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
1
0
0
1
1
0
1
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
1
0
1
0
1
0
1
0
214
Swertia uniflora
Swertia volkensii var. baleensis,var
volkensii
Thymus schimperi
Thymus serrulatus
Trifolium acaule
Trifolium calocephalum
Trifolium cryptopodium
Trifolium elgonense
Trifolium multinerve
Trifolium rueppellianum
Trifolium simense
Umbilicus botryoides
Ursinia nana
Valerianella
microcarpa
var.
microcarpa
Verbascum sedgwickianum
Veronica anagallis-aquatica
Veronica arvensis
Veronica glandulosa
Veronica gunae
Viola eminii
Vulpia bromoides
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
1
0
0
0
0
1
0
0
0
1
0
0
0
0
1
0
0
1
1
0
0
0
0
0
1
0
1
0
0
1
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
1
1
1
0
0
0
0
0
0
0
0
0
1
1
0
0
1
0
0
0
0
0
1
0
0
0
1
0
1
0
1
0
0
0
0
0
1
0
1
0
1
0
0
0
0
0
0
1
1
0
1
0
1
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
1
0
0
0
1
1
1
0
0
0
0
0
1
0
0
0
0
0
0
0
1
1
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
215
Appendix 2a: Data Matrix of the species of Pentaschistis for Phenetic Analysis
Specimens
(OTUs)
ET−1604−8
ET−1604−21
ET−1604−7
ET−1604−1
ET−1604−6
ET−1009−3
ET−717−1
ET−220−3
TZ−116−4
KN−251−4
KN−538−3
TZ−184
KN−251−3
ET−366−1
ET−763−3
ET−902−3
ET−647−3
ET−813−3
A. Bogdan
AB 4123
A. Bogdan
AB 44499
I.R.
Dale
1933
TZ−48
ET−946−3
TZ−453−4
O. Hedberg
4208
O. Hedberg
2301
TZ−454−5
J.M. Thresh
JMT 4
TZ−348−4
CL(cm)
65
120
80
60
75
6
8.5
5.5
11
17
14
13.5
18
21
33
30
28.5
23
LF
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
GLLV
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
LVW
(mm)
6
6
8
6
6
2
3
2
3
2.5
2.5
3
2.5
4
3
3
3
2
PAL(cm)
10
16
13
13
12
3
5
3
8.5
9
7.5
7
8.5
4
6.5
7
6
6
PAW
(cm)
4
3.5
2
4
3
0.3
1
0.7
6
5.5
4
4
4
0.9
1.1
1.2
1.5
1.1
INFL(cm)
8.5
18
8
15
12
1
1
1
4
11
5
8
12.5
5
9
6.5
3.5
4
SPC
0
0
0
0
0
1
1
1
2
2
2
2
2
3
3
3
3
3
No.
SP
200
200
180
180
160
23
80
45
55
65
70
60
60
50
100
90
85
100
SPL
(mm)
8
8
8
9
9
6
6
5
6
6
6
6
6
6
6
7
7
6
GLPA
0
0
0
0
0
1
1
1
1
1
1
1
1
0
0
0
0
0
GLUC
1
1
1
1
1
1
1
4
3
3
3
3
3
3
3
3
3
3
ASGLU
0
0
0
0
0
2
2
1
3
3
3
3
3
0
0
0
0
0
GLUT
1
1
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
SHINF
1
1
1
1
1
0
0
1
1
1
1
1
1
0
0
0
0
0
ANTC
0
0
0
0
0
1
1
3
2
2
2
2
2
0
0
0
0
0
AWL
(mm)
9
12
10
10
10
3
3
6
6
6
7
5.5
6
5
6
7
7
6
APL(mm)
5
6
5
5
5
2
1.2
1
2
2.5
3
2
2
2
2.5
3
2.5
2.5
9
1
0
1
2
0.2
2
2
10
4
0
3
0
0
0
0
3.5
1.5
11
1
0
1
3
0.5
3
2
12
4
0
3
0
0
0
0
4
1.8
17
25
7
25
1
1
1
0
0
0
0
0
3
1
1
2
3.3
6.5
3
6
1.2
0.4
0.3
0.6
4
3
5
9
2
2
2
2
60
17
21
25
6
4
4
6
0
0
0
0
3
3
3
2
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
6
3.5
4
5
3
1.5
2
3
17.5
0
0
3
4.5
1
2
2
30
8
1
2
1
0
0
0
7
3
29
24
0
0
0
0
3
4
5
6.5
1.2
1
1.5
3
2
2
32
30
6
6
1
1
2
2
1
1
0
0
0
0
0
0
8
7
3
3
26
102
0
0
1
0
3
10
4
21
1
4.5
7
8
2
4
35
210
7
10
1
0
2
3
1
1
0
1
0
1
0
0
5
14
2.5
6
216
V. C. Gilbert
2249
Greenway
3801
D.
Vesey
Fitzgerald
4853
D.
Vesey
Fitzgerald
7506
O. Kerfoot
1734
R. Wingfield
350
W.H. Prins
341
P.J. Muller
&J.C.
Scheeper
145
G. Davidse
et. al 6558
107
0
0
10
32
6.5
7
4
220
10
0
3
1
1
1
0
11
5
140
0
0
10
21
5
14.5
4
200
8
0
3
1
1
1
0
10
5
84
0
0
8
19
4.5
10
4
210
9
0
3
1
1
1
0
12
6
87
0
0
8
19
5
10
4
250
10
0
3
1
1
1
0
8
3
55
0
0
4
15
3.5
8
5
150
8
0
5
0
2
1
4
9
5
88
0
0
4
19
5
31
5
100
7
0
5
0
2
1
4
8
3
84
0
0
5
17
14
24
5
120
7
0
5
0
2
1
4
9
4
50
0
0
3
8
3
9
5
34
7
0
5
0
2
1
4
8
3
56
0
0
4
18
8
10
5
120
7
0
5
0
2
1
4
11
5
217
Appendix 2b: Data Matrix
Specimens
(OTUs)
ET−1604−8
ET−1604−21
ET−1604−7
ET−1604−1
ET−1604−6
ET−1009−3
ET−717−1
ET−220−3
TZ−116−4
KN−251−4
KN−538−3
TZ−184
KN−251−3
ET−366−1
ET−763−3
ET−902−3
ET−647−3
ET−813−3
A. Bogdan
AB 4123
A. Bogdan
AB 44499
I.R.
Dale
1933
TZ−48
ET−946−3
TZ−453−4
O. Hedberg
4208
O. Hedberg
2301
TZ−454−5
J.M. Thresh
JMT 4
TZ−348−4
Habit
LEI
LESM
LEL
LEM
GLTYP
GLLOC
LEML
LEMI
GLAWN
PALEAL
PALEI
ANTL
ULPEDL
BPEDL
GLL
CRLINF
FLORL
FLORIND
1
1
1
1
1
1
1
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
1
1
1
1
1
1
1
1
1
1
45
50
45
53
40
3
9
7
12
15
10
9
15
17
15
20
15
20
1
1
1
1
1
1
1
1
0
0
0
0
0
1
1
1
1
1
0
0
0
0
0
2
2
1
1
1
1
1
1
0
0
0
0
0
0
0
0
0
0
2
2
1
1
1
1
1
1
0
0
0
0
0
3
3.5
3
3.5
3
2
2
2
2.2
3
2.5
3
2.5
2
2
2
2
2
1
1
1
1
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
1
1
0
0
0
0
0
4
4.5
4
5
4
2
2
1.5
1.5
3
3.5
3
3
3
2
2.5
2
2
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
2.7
2.6
2.5
2.7
2.6
0.5
0.6
0.7
1
1.5
2
1
1.5
0.5
0.6
0.7
0.5
0.6
4
4
2
4
3
2
2
2
8
7
9
7
7
2
2
2
2
2
4
4
4
4
4
4
4
4
16
16
17
12
16
4
4
4
4
4
8
8
8
9
9
6
6
5
6
6
6
6
6
6
6
7
7
6
0
0
0
0
0
4
4
4
3
3
3
3
3
1
1
1
1
1
4
4
4
4
4
2
2
2
3
2.5
3
2
2.5
2.5
2.5
2.5
3
2.5
2
2
2
2
2
0
0
0
0
0
0
0
0
1
1
1
1
1
1
1
4
1
0
0
2
0
0
2
0
0.5
2
3
4
2
2
1
1
1
6
1
0
0
2
0
0
2
0
0.6
2
3
4
2
2
1
1
1
6
1
0
0
2
0
0
2
0
0.5
2
3
4
2
2
1
1
1
1
1
1
1
7
5
13
1
1
1
0
0
1
0
0
3
2
2
2
0
0
0
0
0
0
2
2
2
0
0
0
0.6
0.5
0.7
2
2
2
3
3
6
4
4
6
2
2
2
2
2
3
1
1
1
1
1
22
1
1
3
2.5
0
0
2.5
0
0.8
2
6
7
2
3
1
1
1
24
1
1
3
2
0
0
2
0
0.9
2
6
6
2
3
1
1
1
15
1
1
3
1.8
0
0
2
0
0.7
2
6
6
2
3
1
1
1
16
1
1
3
2
0
0
2
0
0.7
3
5
7
2
3
1
0
1
56
0
0
0
3
1
0
4.5
0
2.5
4
11
10
1
4
2
1
1
1
1
1
1
1
1
1
1
1
218
V. C. Gilbert
2249
Greenway
3801
D.
Vesey
Fitzgerald
4853
D.
Vesey
Fitzgerald
7506
O. Kerfoot
1734
R. Wingfield
350
W.H. Prins
341
P.J. Muller
&J.C.
Scheeper
145
G. Davidse
et. al 6558
1
1
0
1
45
0
0
0
3.5
1
0
5
0
2.5
5
13
10
1
4
2
0
1
49
0
0
0
3
1
0
4.5
0
2.5
5
12
10
1
4
2
0
1
45
0
0
0
3
1
0
4.5
0
2.3
5
11
10
1
4
2
0
1
43
0
0
0
3.5
1
0
4.5
0
2.5
5
12
10
1
4
2
1
1
28
1
0
0
2
1
0
3.5
2
2
12
17
8
2
4
2
1
1
30
1
0
0
2.5
1
0
4
2
2
10
16
7
2
4
2
1
1
30
1
0
0
3
1
0
3.5
2
2
12
20
7
2
4
2
1
1
30
1
0
0
2.5
1
0
3.5
2
2
12
17
7
2
4
2
1
1
1
1
1
1
219
Appendix 3: Species count for Pentaschistis species from five Vegetation types
Species
Pentaschistis borussica
Pentaschistis pictigluma var gracilis
Pentaschistis pictigluma var. minor
Pentaschistis
pictigluma
var.
pictigluma
Pentaschistis trisetoides
Rock
outcrop
8
0
5
Bog
2
1
1
Grassland
7
0
2
Dendrosenecio
1
0
1
Alchemilla
0
1
0
5
2
4
0
2
0
0
0
0
0
220
Appendix 4: Summary of the Family, genera and species sampled from Eastern Africa
No.
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
Family
Asteraceae
Poaceae
Caryophyllaceae
Lamiaceae
Cyperaceae
Rosaceae
Ranunculaceae
Lobeliaceae
Apiaceae
Scrophulariaceae
Gentianaceae
Fabaceae
Crassulaceae
Brassicaceae
Rubiaceae
Ericaceae
Adiantaceae
Iridaceae
Clusiaceae
Boraginaceae
Aspleniaceae
Asphodelaceae
Juncaceae
Geraniaceae
Dryopteridaceae
Dipsacaceae
Commelinaceae
Callitrichaceae
Woodsiaceae
Violaceae
Valerianalaceae
Urticaceae
Saxifragaceae
Primulaceae
Polygonaceae
Polygalaceae
Plantaginaceae
Papaveraceae
Oxalidaceae
Orchidaceae
No. of
genera
21
17
9
4
4
2
3
1
7
5
1
2
5
5
1
1
3
3
1
2
1
1
1
1
1
2
1
1
1
1
1
1
1
1
1
1
1
1
1
1
Percentage
(%)
16.9
13.7
7.3
3.2
3.2
1.6
2.4
0.8
5.6
4.0
0.8
1.6
4.0
4.0
0.8
0.8
2.4
2.4
0.8
1.6
0.8
0.8
0.8
0.8
0.8
1.6
0.8
0.8
0.8
0.8
0.8
0.8
0.8
0.8
0.8
0.8
0.8
0.8
0.8
0.8
No. of
Species
69
37
16
12
11
10
9
9
9
8
8
8
8
6
4
4
4
3
3
3
3
3
2
2
2
2
2
2
1
1
1
1
1
1
1
1
1
1
2
1
Percentage
(%)
24.8
13.3
5.8
4.3
3.9
3.6
3.2
3.2
3.2
2.9
2.9
2.9
2.9
2.2
1.4
1.4
1.4
1.0
1.0
1.0
1.0
1.0
0.7
0.7
0.7
0.7
0.7
0.4
0.4
0.4
0.4
0.4
0.4
0.4
0.4
0.4
0.4
0.7
0.4
221
41
42
43
44
45
46
Onagraceae
Malvaceae
Lycopodiaceae
Eriocaulaceae
Colchicaceae
Campanulaceae
1
1
1
1
1
1
0.8
0.8
0.8
0.8
0.8
0.8
1
1
1
1
1
1
0.4
0.4
0.4
0.4
0.4
0.4
KK= Kenya Mt. Kenya; KE = Kenya Elgon; TK =Tanzania, Kilimanjaro; UR =Uganda,
Ruwenzori; EB =Ethiopia, Bale; ES =Ethiopia, Simen; R= rock outcrop, B=bog, G=
grassland, D =Dendrosenecio, A =Alchemilla.
222
Appendix 5: Comparison of Species Diversity & richness among the 75 plots from Eastern
African mountains. The green colour indicates the most important factors that distinguish
taxa
Plot
No.
KKRP1
KKRP2
KKRP3
Species
Count
28
30
23
Margale'f
(d)
8.10274397
8.52640901
7.016437756
Brillouin
2.424634
2.488608
2.243769
H'(loge)
3.332205
3.401197
3.135494
KKRP4
KKBP1
KKBP2
KKBP3
KKBP4
KKDP1
KKDP2
KKDP3
KKDP4
KERP1
KERP2
KERP3
KERP4
KEGP1
KEGP2
KEGP3
KEGP4
KEDP1
KEDP2
KEDP3
TKRP1
TKRP2
TKRP3
TKRP4
TKBP1
TKBP2
TKBP3
TKBP4
TKGP1
TKGP2
TKGP3
TKGP4
URBP1
URBP2
URBP3
17
23
24
20
41
19
22
17
16
33
32
35
26
32
21
24
35
22
22
22
25
21
35
20
38
31
21
20
23
23
14
14
43
28
15
5.647297982
7.016437756
7.23713355
6.342355813
10.77130032
6.113218894
6.793824516
5.647297982
5.410106403
9.15198936
8.944709254
9.563058078
7.673191911
8.944709254
6.569174775
7.23713355
9.563058078
6.793824516
6.793824516
6.793824516
7.456019215
6.569174775
9.563058078
6.342355813
10.17158038
8.736200287
6.569174775
6.342355813
7.016437756
7.016437756
4.926001362
4.926001362
11.1666486
8.10274397
5.169771223
1.970887
2.243769
2.282697
2.116781
2.781322
2.07052
2.203236
1.970887
1.916991
2.577408
2.548686
2.632462
2.356219
2.548686
2.160959
2.282697
2.632462
2.203236
2.203236
2.203236
2.320144
2.160959
2.632462
2.116781
2.709689
2.519104
2.160959
2.116781
2.243769
2.243769
1.799373
1.799373
2.826351
2.424634
1.859951
2.833213
3.135494
3.178054
2.995732
3.713572
2.944439
3.091042
2.833213
2.772589
3.496508
3.465736
3.555348
3.258097
3.465736
3.044522
3.178054
3.555348
3.091042
3.091042
3.091042
3.218876
3.044522
3.555348
2.995732
3.637586
3.433987
3.044522
2.995732
3.135494
3.135494
2.639057
2.639057
3.7612
3.332205
2.70805
223
URBP4
URDP1
URDP2
URDP3
URDP4
URAP1
URAP2
URAP3
URAP4
EBRP1
EBRP2
EBRP3
EBRP4
EBBP1
EBBP2
EBBP3
EBBP4
EBGP1
EBGP2
EBGP3
EBGP4
EBAP1
EBAP2
EBAP3
EBAP4
ESRP1
ESRP2
ESRP3
ESRP4
ESBP1
ESBP2
ESBP3
ESBP4
ESGP1
ESGP2
ESGP3
ESGP4
12
20
18
10
15
20
22
14
22
28
34
42
32
21
39
25
29
30
20
19
20
27
19
16
20
57
51
43
48
44
37
40
34
45
40
45
31
4.426725648
6.342355813
5.881596356
3.908650337
5.169771223
6.342355813
6.793824516
4.926001362
6.793824516
8.10274397
9.358090238
10.96940184
8.944709254
6.569174775
10.37241998
7.456019215
8.315277722
8.52640901
6.342355813
6.113218894
6.342355813
7.888739964
6.113218894
5.410106403
6.342355813
13.85092503
12.71673891
11.1666486
12.14093504
11.36306691
9.969764163
10.5723162
9.358090238
11.55868168
10.5723162
11.55868168
8.736200287
1.665601
2.116781
2.021969
1.510441
1.859951
2.116781
2.203236
1.799373
2.203236
2.424634
2.605318
2.804092
2.548686
2.160959
2.734148
2.320144
2.457139
2.488608
2.116781
2.07052
2.116781
2.39102
2.07052
1.916991
2.116781
3.094664
2.988423
2.826351
2.930707
2.84812
2.684611
2.758016
2.605318
2.869421
2.758016
2.869421
2.519104
2.484907
2.995732
2.890372
2.302585
2.70805
2.995732
3.091042
2.639057
3.091042
3.332205
3.526361
3.73767
3.465736
3.044522
3.663562
3.218876
3.367296
3.401197
2.995732
2.944439
2.995732
3.295837
2.944439
2.772589
2.995732
4.043051
3.931826
3.7612
3.871201
3.78419
3.610918
3.688879
3.526361
3.806662
3.688879
3.806662
3.433987
224
Appendix 6: Factor coordinates of the variables, based on correlations (α=0.05) showing
contribution of each character
Variables
Factor 1
Factor 2
Culm length (cm)
−0.931808
0.139898
Leaf form
0.402137
0.335374
Glands on leaf
0.341640
−0.061741
Leaf width (mm)
−0.879374
0.140551
Panicle length(cm)
−0.922147
−0.150105
Panicle width (cm)
−0.675553
−0.554726
Inflorescence stalk length (cm)
−0.653765
−0.238590
Spikelet colour
−0.359071
−0.362657
Number of spikelets
−0.921263
0.139919
Spikelet length (mm)
−0.847205
0.092779
Glands on panicle
0.518975
−0.550953
Glume colour
−0.142005
−0.526447
Apex shape of glume
0.240128
−0.703352
Glume indumentum
−0.846531
−0.169860
Shape of inflorescence
−0.743564
−0.477475
Anther colour
−0.123738
−0.830395
Awn length (mm)
−0.906432
0.052024
Apical bristle length (mm)
−0.883746
0.184985
Habit
−0.189322
0.149026
Leaf Indumentum
0.615114
−0.051794
Leaf sheath mouth
−0.189322
0.149026
Leaf length
−0.931239
0.221056
Leaf margin
0.333003
0.497351
Gland type
0.565486
−0.455049
Gland location
0.498750
−0.182795
Lemma length
−0.808909
−0.058095
Lemma indumentum
−0.881843
0.001280
Presence of glands on awn
0.182530
−0.697871
Palea length
−0.928655
0.012457
Palea indumentum
−0.244850
−0.471305
Anther length
−0.926487
−0.075016
Ultimate pedicel length
−0.477449
−0.770043
Basal Pedicel length
−0.454288
−0.843252
225
Glume length
−0.875207
0.059071
Colour of Inflorescence
0.628223
−0.682671
Floret length
−0.929760
0.006535
Floret indumentum
−0.832789
0.410002
226
Appendix 7: Data matrix for species richness of each vegetation community
Vegetation Community/Plot No.
ROCK
BOG
GRASSLAND
DENDROSENECIO
ALCHEMILLA
1
28
23
32
19
20
2
30
24
21
22
22
3
23
20
24
17
14
4
17
41
35
16
22
5
33
43
23
22
27
6
32
28
23
22
19
7
35
15
14
22
16
8
26
12
14
20
20
9
25
38
30
18
10
21
31
20
10
15
11
35
21
19
12
20
20
20
13
28
21
45
14
34
39
40
15
42
25
45
16
32
29
31
17
57
44
18
51
37
19
43
40
20
48
34
227
Appendix 8: Data matrix for species richness of each East African Mountain
Mountain/Plot
No.
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
Kenya
28
30
23
17
23
24
20
41
19
22
17
16
Elgon
33
32
35
26
32
21
24
35
22
22
22
Kilimanjaro
25
21
35
20
38
31
21
20
23
23
14
14
Rwenzori
43
28
15
12
20
18
10
15
20
22
14
22
Bale
28
34
42
32
21
39
25
29
30
20
19
20
27
19
16
20
Simen
57
51
43
48
44
37
40
44
45
40
45
31
228
Appendix 9: Species checklist of Afroalpine Flora from Eastern Africa
Family
Adiantaceae
Adiantaceae
Apiaceae
Apiaceae
Apiaceae
Apiaceae
Apiaceae
Apiaceae
Apiaceae
Araliaceae
Asphodelaceae
Asphodelaceae
Asphodelaceae
Aspleniaceae
Aspleniaceae
Aspleniaceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Species
Adiantum thalictroides Willd. ex Schlechtend.
Cheilanthes farinosa (Forssk.) Kaulf.
Agrocharis melanantha Hochst.
Heracleum abyssincum (Boiss.) Norman
Heracleum elgonense (H.Wolff) Bullock
Peucedanum kerstenii Engl.
Pimpinella oreophila Hook.f.
Pimpinella pimpinelloides H.Wolff
Haplosciadium abyssinicum Hochst.
Hydrocotylee sibthorpioides Lam.
Kniphofia foliosa Hochst.
Kniphofia isoetifolia A.Rich.
Kniphofia thomsonii Baker
Asplenium abyssinicum Fée
Asplenium aethiopicum (Burm.f.) Bech
Asplenium buttneri Hook.
Helichrysum citrispinum Delile
Helichrysum foetidum (L.) Cass.
Helichrysum formosissimum Sch.Bip.
Helichrysum forskahlii (J.F.Gmel.) Hilliard & B.L.Burtt
Helichrysum globosum Sch.Bip.
Helichrysum gofense Cufod.
Helichrysum horridum Sch.Bip.
Helichrysum kilimanjari Oliv.
Helichrysum newii Oliv. & Hiern
Helichrysum ambylyphyllum Mattf.
Helichrysum argyranthum O.Hoffm.
Helichrysum splendidum (Thunb.) Less.
Helichrysum stuhlmannii O.Hoffm.
Anthemis tigrensis A.Rich.
Artemisia afra Jacq. ex Willd.
Carduus chamaecephalus (Vatke) Oliv. & Hiern
Carduus keniensis R.E.Fr.
Carduus leptacanthus Fresen.
Carduus macracanthus Kazmi
Carduus ruwenzoriensis (Cortesi) Chiov.
Carduus schimperi Sch. Bip.
Cineraria abyssinica Sch.Bip. ex A.Rich.
Cineraria deltoidea Sond.
229
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Conyza spinosa Sch.Bip. ex Oliv. & Hiern
Conyza subscaposa O.Hoffm.
Cotula abyssinica Sch.Bip.
Cotula cryptocephala Sch.Bip. ex A.Rich
Crepis foetida L.
Crepis rueppellii Sch.Bip.
Dendrosenecio adnivalis (Stapf) E.B.Knox ssp. adnivalis var. adnivalis
Dendrosenecio battiscombei (R.E.Fr. & T.C.E.Fr.) E.B.Knox
Dendrosenecio elgonensis (T.C.E.Fr.) E.B.Knox ssp. barbatipes
Dendrosenecio erici-rosenii (R.E.Fr. & T.C.E.Fr.) E.B.Knox ssp. erici-rosenii
Dendrosenecio keniensis (Baker f.) Mabb.
Dendrosenecio keniensis × D. keniodendron
Dendrosenecio keniodendron (r.e.fr. & t
Dendrosenecio kilimanjari (Mildbr.) E.B.Knox ssp. cottonii
Dichrocephala chrysanthemifolia (Blume) DC.
Echinops buhaitensis Mesfin
Erigeron alpinus L.
Euryops dacrydioides Oliv.
Euryops elgonensis Mattf.
Euryops prostratus B.Nord.
Gnaphalium unionis Sch.Bip. ex Oliv. & Hiern
Haplocarpha rueppellii (Sch.Bip.) Beauverd
Haplocarpha schimperi (Sch.Bip.) Beauverd
Helichrysum brownei S.Moore var. brownei
Lactuca inermis Forssk.
Oreophyton falcatum (A.Rich.) O.E.Schulz
Phagnalon abyssinicum Sch.Bip. ex Hochst.
Senecio balensis S.Ortiz & Vivero
Senecio farinaceous Sch.Bip. ex A.Rich.
Senecio fresenii Sch.Bip.
Senecio jacksonii S. Moore
Senecio meyeri-johannis Engl.
Senecio nanus Sch.Bip. ex A.Rich.
Senecio polyadenus Hedberg
Senecio purtschelleri Engl.
Senecio rhammatophyllus Mattf.
Senecio schultzii Wedd.
Senecio schweinfurthii O. Hoffm.
Senecio subsessilis Oliv. & Hiern
Senecio telekii (Schweinf.) O.Hoffm.
Senecio transmarinus S. Moore
Senecio transmarinus S. Moore var. major
230
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Athyriaceae
Athyriaceae
Athyriaceae
Boraginaceae
Boraginaceae
Brassicaceae
Brassicaceae
Brassicaceae
Brassicaceae
Brassicaceae
Brassicaceae
Brassicaceae
Callitrichaceae
Callitrichaceae
Campanulaceae
Caryophylaceae
Caryophylaceae
Caryophylaceae
Caryophylaceae
Caryophylaceae
Caryophylaceae
Caryophylaceae
Caryophylaceae
Caryophylaceae
Caryophylaceae
Caryophylaceae
Caryophylaceae
Caryophylaceae
Caryophylaceae
Colchicaceae
Commelinaceae
Commelinaceae
Crassulaceae
Crassulaceae
Crassulaceae
Crassulaceae
Senecio transmarinus S. Moore var. sycephyllus
Senecio unionis Sch.Bip. ex A.Rich.
Sonchus melanolepis Fresen.
Stoebe kilimandscharica O.Hoffm.
Ursinia nana DC
Crepis dianthoseris N.Kilian, Enke, Sileshi & Gemeinholzer
Cystopteris fragilis (L.) Bernh.
Polystichum magnificum F.Ballard
Polystichum setiferum (Forssk.) Moore ex Woynar
Myosotis keniensis T.C.E.Fr.
Myosotis vestergrenii Stroh
Arabidopsis thaliana (L.) Heynh.
Arabis alpina L.
Barbarea intermedia Boreau
Cardamine hirsuta L.
Cardamine obliqua A.Rich.
Erophila verna (L.) Chevall.
Subularia monticola Schweinf.
Callitriche oreophila Schotsman
Callitriche vulcanicola Schotsman
Campanula edulis Forssk.
Cerastium afromontanum T.C.E.Fr. & Weim
Cerastium octandrum Hochst. ex A.Rich.
Dianthus longiglumis Delile
Herniaria abyssinica Chaudhri
Lychnis abyssinica (Hochst.) Lidén
Lychnis rotundifolia (Oliv.) M.Popp
Minuartia filifolia (Forssk.) Mattf.
Paronychia bryoides Hochst. ex A.Rich.
Sagina abyssinica [Hochst ex] A. Rich.
Sagina afroalpina Hedberg
Silene burchellii Otth ex DC. var burchellii
Silene flammulifolia Steud. ex A. Rich.
Silene macrosolen Steud. ex A.Rich.
Stellaria media (L.) Vill.
Merendera schimperiana Hochst.
Cyanotis polyrrhiza Hochst. ex Hassk.
Cyanotis barbata D.Don
Crassula granvikii Mildbr.
Crassula schimperi Fisch. & C.A.Mey.
Rosularia semiensis (J. Gay ex A. Richard) H. Ohba
Sedum crassularia Raym.-Hamet
231
Crassulaceae
Crassulaceae
Crassulaceae
Crassulaceae
Cyperaceae
Cyperaceae
Cyperaceae
Cyperaceae
Cyperaceae
Cyperaceae
Cyperaceae
Cyperaceae
Cyperaceae
Cyperaceae
Cyperaceae
Dipsacaceae
Dipsacaceae
Ericaceae
Ericaceae
Ericaceae
Ericaceae
Eriocaulaceae
Fabaceae
Fabaceae
Fabaceae
Fabaceae
Fabaceae
Fabaceae
Fabaceae
Fabaceae
Gentianaceae
Gentianaceae
Gentianaceae
Gentianaceae
Gentianaceae
Gentianaceae
Gentianaceae
Gentianaceae
Gentianaceae
Gentianaceae
Hypericaceae
Hypericaceae
Sedum meyeri-johannis Engl.
Sedum mooneyi M.G.Gilbert
Umbilicus botryoides Hochst. ex A.Rich.
Aeonium leucoblepharum Webb ex A. Richard
Carex monostachya A.Rich.
Carex runssoroensis var. aberdarensis K.Schum.
Carex simensis A.Rich.
Carex conferta A.Rich.
Carpha eminii (K.Schum.) C.B.Clarke.
Carex bequaerttii De Wild.
Cyperus elegantulus Steud.
Cyperus plateilema (Steud.) Kuk.
Isolepis costata Hochst. ex A.Rich.
Isolepis fluitans (L.) R.Br. var. fluitans, var. major
Isolepis setacea L.) R.Br.
Dipsacus pinnatifidus A.Rich.
Scabiosa columbaria L
Erica arborea L.
Erica johnstonii (Schweinf. ex Engl.) Dorr
Erica tenuipilosa (Engl.) R.E. Fries ssp. spicata
Erica trimera (Engl.) R.E. Fries ssp. keniensis
Eriocaulon schimperi Körn. ex Ruhland.
Argyrolobium schimperianum Hochst.
Trifolium acaule A. Rich.
Trifolium calocephalum Fresen.
Trifolium cryptopodium A.Rich.
Trifolium elgonense Gillett
Trifolium multinerve A.Rich.
Trifolium rueppellianum Fresen.
Trifolium simense Fresen.
Swertia abyssinica Hochst.
Swertia crassiuscula Gilg. ssp. crassiuscula-var. cras & leuc;ssp. robusta
Swertia engleri var. engleri (ssp. Engleri,ssp. woodii)
Swertia kilimandscharica Engl.
Swertia pumila Hochst. ex Hook.f.
Swertia subnivalis T.C.E. Fr.
Swertia uniflora Mildbr.
Swertia volkensii Gilg. var. baleensis,var volkensii
Geranium arabicum Forssk.
Geranium sp.nov.=Miehe 3002
Hypericum peplidifolium A.Rich.
Hypericum revolutum Vahl
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Hypericaceae
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Juncaceae
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Hypericum afromontanum Bulloc
Dierama cupuliflorum Klatt
Hesperantha petitiana (A.Rich.) Baker
Limosella major Diels
Romulea fischeri Pax
Luzula abyssinica Parl.
Luzula johnstonii Parl.
Salvia nilotica Jacq.
Satureja abyssinica (Benth.) Briq.
Satureja imbricata Briq.
Satureja kilimandschari (Gurke) Ryding
Satureja pseudosimensis Brenan
Satureja punctata (Benth.) Briq.ssp. punctata
Satureja simensis Briq.
Satureja uhligii (Gurke) Ryding var. obtusifolium
Thymus serrulatus Hochst. ex Delile
Thymus schimperi Ronniger
Lobelia aberdarica R.E.Fr. & T.C.E.Fr.
Lobelia bequaertii De Wild.
Lobelia deckenii (Ascher) Hemsl. ssp. deckenii, ssp. burttii
Lobelia telekii ssp. elgonensis, gregororiana
Lobelia lindblomii Mildbr.
Lobelia rhynchopetalum
Lobelia stuhlmannii Stuhlmann
Lobelia telekii Schweinf.
Lobelia wollastonii Bak f.
Huperzia saururus (Lam.) Trevis.
Malva verticillata L.
Epilobium stereophyllum Fresen. var. stereophyllum
Disa stairsii Kraenzl.
Oxalis corniculata L
Oxalis obliquifolia A.Rich.
Plantago afra L. var. stricta
Agrostis quinqueseta (Steud.) Hochst
Agrostis schimperiana Steud.
Agrostis sclerophylla C.E.Hubb.
Agrostis trachyphylla Pilg.
Agrostis volkensii Stapf
Aira caryophyllea L.
Alopecurus baptarrhenius S.M.Phillips
Andropogon amethystinus Steud.
Andropogon lima (Hack.) Stapf
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Anthoxanthum nivale K.Schum.
Bromus leptoclados Nees
Colpodium chionogeiton (Pilg.) Tzvelev
Colpodium hedbergii (Melderis) Tzvelev
Deschampsia angusta Stapf & C.E.Hubb.
Deschampsia caespitosa (L.) P.Beauv.
Deschampsia flexuosa var. afromontana (L.) Trin.
Festuca abyssinica Hochst. ex A.Rich.
Festuca macrophylla E.B.Alexeev
Festuca pilgeri St. Yves
Festuca richardii E.B.Alexeev
Festuca simensis A.Rich.
Helictotrichon elongatum (A.Rich.) C.E.Hubb.
Helictotrichon cf umbrosum (Steud.) C.E.Hubb.
Koeleria capensis (Steud.) Nees
Pennisetum humile Hochst. ex A.Rich.
Pentaschistis borussica (K. Schum.) Pilg.
Pentaschistis pictigluma (Steud.) Pilg. var gracilis
Pentaschistis pictigluma (Steud.) Pilg. var. minor
Pentaschistis pictigluma (Steud.) Pilg. var. pictigluma
Pentaschistis trisetoides Pilg.
Poa annua L.
Poa leptoclada A.Rich.
Poa schimperiana Hochst. ex A.Rich.
Polypogon schimperianus (Hochst. ex Steud.) Cope)
Rytidosperma grandiflorum (Hochst. ex A.Rich.) S.M.Phillips
Rytidosperma subulata (A.Rich.) T.A.Cope
Vulpia bromoides (L.) Gray
Agrostis gracilifolia C.E.Hubb ssp. gracilifolia S.M. Philips
Polygala steudneri Chodat
Rumex nepalensis Spreng.
Anagallis serpens ssp. meyeri-johannis (Engl.) P.Taylor
Anogramma leptophylla (L.) Link
Delphinium macrocentrum Oliv.
Delphinium wellbyi Hemsl.
Ranunculus distrias Steud. ex A.Rich.
Ranunculus oligocarpus Speg.
Ranunculus oreophytus Del.
Ranunculus stagnalis Hochst. ex A.Rich.
Ranunculus trichophyllus Chaix
Ranunculus volkensii Engl.
Alchemilla abyssinicaFresen.
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Alchemilla argyrophylla ssp. argyrophylla Oliv.
Alchemilla haumannii Engl.
Alchemilla johnstonii Oliv.
Alchemilla microbetula T.C.E. Fries
Alchemilla pedata A. Rich.
Alchemilla subnivalis Baker f.
Alchemilla stuhlmannii Engl.
Alchemilla triphylla Rothm.
Anemone thomsonii Oliv. var. thomsonii
Aphanes bachitii (Hauman & Balle) Rotm.
Alchemilla argyrophylla Oliv. ssp. argyrophylloides (Baker f.) Rothm
Alchemilla argyrophylla Oliv. ssp. argyrophylla Oliv.
Galium acrophyumHochst. ex Chiov.
Galium ruwenzoriense (Cortesi) Chiov.
Galium simense Fresen.
Galium ossirwaense K.Krause var. glabrum
Galium thunbergianum Eckl. & Zeyh.
Saxifraga hederifolia Hochst. ex A.Rich.
Bartsia longiflora ssp. longiflora Hochst. ex Benth.
Hebenstretia angolensis Rolfe
Lithospermum afromontanum Weim
Rhabdotosperma scrophulariifolia (Hochst.) Hartl
Verbascum sedgwickianum (Schimp.) Hub.
Veronica anagallis-aquatica L.
Veronica arvensis L.
Veronica glandulosa Hochst ex. Benth.
Veronica gunae Schweinf. ex Engl.
Bartsia decurva Hochst. ex Benth.
Parietaria debilis G.Forst.
Valerianella microcarpa Loisel. var microcarpa
Viola eminii (Engl.) R.E.Fr.
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Appendix 10: Plates of Some typical Afroalpine species from Eastern Africa….
Plate 1 Carduus keniensis
(Saddle hut, Mt. Meru taken on 27/11/08)
Plate 3 Cardamine obliqua
(Bujuku, Rwenzori on 28/8/08)
Plate 2 Anemone thomsonii
(Shira, Mt. Kilimanjaro on 4/11/08)
Plate 4 Alchemilla johnstonii
(Mt Meru, near Crater on 30/11/08)
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Plate 5 Dendrosenecio advinalis var. petiolatus
(Bujuku, Rwenzori on 21/08/08)
Plate 7 Galium ruwenzoriense
(Bujuku, Rwenzori taken on 24/8/08)
Plate 6 Disa stairsii
(John Mate hut, Rwenzori on 14/8/08)
Plate 8 Haplocarpha rueppellii
(Baranko, Mt Kilimanjaro on 8/11/08)
237
Plate 9 Gladiolus watsonioides
(Near Saddle hut Mt Meru on 30/11/08)
Plate 11 Helichrysum formosissimum
(Baranko hut, Mt Kilimanjaro on 8/11/08)
Plate 10 Ranunculus oreophytus
(Lower Bukurungu, Rwenzori on 17/8/08)
Plate 12 Hebenestreitia angolensis
(Near Saddle hut Mt Meru on 30/11/08)
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Plate 15 Subularia monticola
Plate 13 Galium acrophyum
(Saddle hut Mt Meru on 30/11/08)
Plate 15: Subularia monticola
(Bujuku, Rwenzori on 28/8/08)
Plate 16 Huperzia saururus
Plate 14 Helichrysum forskahlii
(Baranko, Mt Kilimanjaro on 8/11/08)
Plate 16: Huperzia saururus
(Bujuku, Rwenzori on 20/8/08)
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Appendix 11: Botanical illustrations of Pentaschistis species
Illustration: 17 (a)– (f): (a) Pentaschistis pictigluma var. minor– growth habit; (b) P.
chrysurus– (i) Inflorescences ii) a section of the stem and leaf sheath; (c) P. trisetoides–
habit; (d) P. borussica (i) habit, (ii) inflorescence; (e) P. natalensis, (i) inflorescence, (ii) leaf
and leaf sheath; (f) P. gracilis var. gracilis– growth habit (Botanical illustration by Nicholus
M.)
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Abdikadir Ahmed