Quaternary International 425 (2016) 301e386

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Quaternary International
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Atlas of pollen and spores and their parent taxa of Mt Kilimanjaro and
tropical East Africa
Lisa Schüler a, *, Andreas Hemp b
a €ttingen University, Untere Karspüle 2, 37073
Department of Palynology and Climate Dynamics, Albrecht-von-Haller Institute for Plant Sciences, Go
Go€ttingen, Germany
b €tsstr. 30, 95440 Bayreuth, Germany
Department of Plant Systematics, Bayreuth University, Universita

a r t i c l e i n f o a b s t r a c t

Article history: The accurate and consistent identification of fossil pollen is essential to allow robust inferences to be
Received 9 May 2016 drawn with regard to past vegetation and climate change. Identifications are best achieved through the
Received in revised form direct inspection of reference material. Most substantial reference collections are located at research
19 July 2016
institutions in Europe or North America, which restricts access for researchers trying to advance paly-
Accepted 25 July 2016
Available online 17 October 2016
nology in less developed countries. Due to the development of digital imaging and fast internet access it
is now possible to produce high quality images from pollen and spore reference collections and make
them globally available. In this pollen and spore atlas we contribute to this growing body of work by
Keywords:
Pollen and spore morphology
presenting images of 240 pollen types (in 202 genera and ~79 families) and 30 spore types (in 25 genera
Palaeoecology and ~17 families) from a wide range of different vegetation types originating from the Kilimanjaro area
Distribution and tropical East Africa. We provide an overview on the range of pollen and spore types commonly found
Ecology in Last Glacial and Holocene sedimentary archives in studies from the Kilimanjaro area and tropical East
Vegetation reconstruction Africa. Besides a pollen key, detailed information is given on pollination, habitus and habitat which all
support the interpretation and environmental reconstructions from pollen records. The atlas will serve as
useful guide for palynological investigations, aiming at detailed and comprehensive reconstruction of
past vegetation, environmental and climate change in tropical East Africa.
© 2016 Elsevier Ltd and INQUA. All rights reserved.

1. Introduction Palynologist can provide a variety of information on and evi-

Our understanding of past vegetation and environmental con-
ditions comes mainly from the examination of macrofossils (e.g.
seeds and wood) and microfossils (e.g. pollen and spores) found in
sedimentary cores. Pollen analysis has been used to document
long-term vegetation dynamics on a landscape scale ever since the
success of Von Post's pioneering experiments beginning in the early
19th century (Von Post, 1916, 1946) and has been subsequently
used to understand changes in vegetation and biodiversity patterns
as well as to address conservation issues (Willis et al., 2007). The
analysis of fossil pollen and spores is now widely used to answer
ecological questions linking past vegetation and environmental
change as well as its response to the occurrence of fires, human
impact and animal activity.
* Corresponding author.
E-mail addresses: lisa.schueler@biologie.uni-goettingen.de (L. Schüler), andreas.
hemp@uni-bayreuth.de (A. Hemp).
dence for past changes in biodiversity (e.g. Odgaard, 1999; Willis
et al., 2007, 2010), ecosystems and biomes (e.g. Prentice et al.,
1996; Jolly et al., 1998; Elenga et al., 2000; Marchant et al., 2009)
as well as climate (Guiot et al., 1989; Chevalier and Chase, 2016;
Marret et al., 2006) but the reliability and dependability of these
information is based entirely on correct and consistent identifica-
tion of the pollen grains and spores in the pollen samples. Hence,
the accurate identification of pollen and spores is crucial for the
reconstruction of past vegetation and subsequent deduction and
interpretation of environmental and climate change.
The number of available pollen and/or spore atlases is increasing
and facilitates the international standardization of identifications
(see also Hooghiemstra and van Geel, 1998). Several print atlases
have already been published for African pollen and spore identifi-
cation (Van Zinderen Bakker, 1953; Maley, 1970; Bonnefille, 1971;
Sowunmi, 1973; Assemien et al., 1974; Riollet and Bonnefille,
1976; Ybert, 1979; Bonnefille and Riollet, 1980; Scott, 1982; Van
Zinderen Bakker and Coetzee, 1988; El Ghazali, Gamal E. B, 1993;
Sowunmi, 1995; Reille, 1999; Gosling et al., 2013) as well as one

http://dx.doi.org/10.1016/j.quaint.2016.07.038
1040-6182/© 2016 Elsevier Ltd and INQUA. All rights reserved.
302 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
online African pollen database (Le zine, 2001; Vincens et al., 2007).
All of them contain hundreds to thousands of images and de-
scriptions of species and therefore provide a good general coverage
of taxa likely to be found in the fossil record or surface samples and
pollen traps. Only the atlas by Riollet and Bonnefille (1976) spe-
cifically covers pollen grains (only Amaranthaceae) from East Af-
rican vegetation types. A compilation of the most common and
ecologically important pollen and spore types from East Africa is
still missing and badly needed.
With this pollen and spore atlas for tropical East Africa we fill
this gap and contribute 715 images and ecological background for
240 pollen and 30 spore taxa commonly found in sediment cores in
this area. Since detailed knowledge on pollen production and
dispersal is important when it comes to the quantitative analysis
and interpretation of pollen and spore data (e.g. Regal, 1982;
Ackerman, 2000; Duffin and Bunting, 2008; Bunting and
Middleton, 2009; Jantz et al., 2013), we included information on
reproduction and pollination agents for each plant family and
species. A short description of the species habitus and habitat is
given to simplify the conceivability of vegetation types from pollen
records. The more general information on habitus, habitat, polli-
nation and pollen morphology given for the different families
makes this atlas applicable for areas where the specific species
described in this atlas do not occur but other species or genera of
this family do.
Generally, the taxa presented in this tropical East African pollen
atlas were selected based upon their (i) common occurence within
fossil pollen records obtained within the region (e.g. Kiage and Liu,
2006; Mumbi et al., 2008; Finch et al., 2009; Rucina et al., 2009;
Zech et al., 2011; Schüler et al., 2012, 2014; Schüler, 2013;
Githumbi et al., 2016) and (ii) they are identified as significant in
the regional flora (Hemp, 2002, 2006a, 2006c; Lovett and Wasser,
2008).
Further, several species of the same genus (e.g. Combretum,
Impatiens, Euphorbia, Indigofera, Podocarpus, Psychotria, Gnidia and
others) are included in this atlas in order to advise caution in the
interpretation of pollen types which may include several species of
one genera. Despite their morphologically similar pollen types
within a genus, the species may occur in different habitats and have
different environmental requirements which needs to be consid-
ered when interpreting their occurrence and abundance in pollen
archives. This matter is especially relevant for highly biodiverse
ecosystems such as the Tropics.
2. Regional setting
2.1. Geology and geography
The East African Rift System is an active continental rift zone
that appears to be a developing divergent tectonic plate boundary
(Baker et al., 1972). The rift is a narrow zone in which the African
Plate is in the process of splitting into two tectonic sub-plates called
the Somali Plate and the Nubian Plate.
Mt Kilimanjaro (3
049 S, 37
219 E) is located in the highlands of
equatorial East Africa in north-eastern Tanzania at the border to
Kenya. It was formed about 3 million years ago during the forma-
tion of the Great Rift Valley (Frisch et al., 2010). About 1 million
years ago the volcanic activities centred on the three points, Shira,
Mawenzi and Kibo (Downie et al., 1956). Mt Kilimanjaro is not only
the highest mountain in Africa but also the highest free-standing
mountain in the world overtopping its surrounding by more than
5000 m (Kaser et al., 2004). The massive is of strato-volcanic origin
and has a diameter of 80 by 50 km. Mt Kilimanjaro is located
roughly 370 km south of the equator and of the Indian Ocean
located to the east.
2.2. Climate
The climate system in tropical Africa responds to a variety of
ocean-atmosphere processes associated with both the Indian and
the Atlantic Ocean (Camberlin et al., 2001). A mixture of Atlantic
and Indian Ocean related mechanisms account for trends in mois-
ture balance in the central and eastern African tropics. In the
easternmost part of tropical Africa, the hydroclimate is oreo-
graphically insulated from the influence of the Atlantic Ocean by
the Congo Air Boundary (CAB) (Sepulchre et al., 2006). Therefore,
this region is primarily influenced by the Indo-Pacific climate
system.
The Kilimanjaro area is characterized by a seasonally dry, trop-
ical climate (Müller, 1983). Due to the ITCZ (Inter-Tropical Conver-
gence Zone) the area further experiences by a bimodal rainfall
pattern with four hydrological seasons per year: two dry and two
wet seasons. The climate on Mt Kilimanjaro is also dependent on
the elevation on the mountain and the exposition of the moun-
tainside. The foothills of the southern slopes receive a mean annual
precipitation (MAP) of 800e900 mm and the lower slopes at
1500 m receive 1500e2000 mm. The forest belt between 2000 and
2300 m receives partly over 3000 mm (Hemp, 2001a). In the alpine
zone precipitation decreases to 200 mm. Due to the prevalence of
SE trade winds, the southern slope is generally wetter than the
northern slope (Hemp, 2006a). The mean annual temperature
(MAT) is 24
C at the southern foothills at about 800 m asl (above
sea level) and decreases linearly by 0.56
C per 100 m altitude
difference to 7.1
C at 5895 m asl (Walter et al., 1975; Thompson
et al., 2002). Frosts can occur above 2700 m asl.
3. Modern vegetation: the source for pollen and spores
3.1. Modern vegetation in East Africa
3.1.1. Coastal vegetation
The vegetation of the coastal area is confined to a narrow strip of
5e10 km with an annual rainfall of about 1000e1400 mm. Sand
dunes, coastal bushland and forest (evergreen lowland forest in
Fig. 1), coastal savanna and mangrove swamps are a feature of some
parts of the coast line of Somalia, Kenya and Tanzania.
However, this area is densely settled for many centuries and
therefore characterized by commercial plantations of coconut
palms and sisal, small holder fields with manihot and maize and
degraded, overgrazed grasslands.
3.1.2. Savanna
Stretching from the coast inland savanna vegetation covers large
areas of East Africa, e.g. two third of Kenya, at low to medium el-
evations of 50e1400 m and 250e900 mm precipitation (Lind and
Morrison, 1974; Friis, 1992). This vegetation type comprises grass-
land with a more or less dense shrub and tree layer, closed grass-
land, open grassland with sparse shrubs, sparse to open grassland,
open deciduous shrubland in Fig. 1. This vegetation formation
typical of Africa was shaped by the long lasting influence of humans
(pastoralism, fire) and herbivores.
The semi-desert vegetation of the driest areas of north Kenya
and Somalia receiving only about 250 mm erratic and unpredict-
able rainfall consist of isolated tufts of grass (Aristida) and occa-
sional low, partly thorny bushes (Commiphora, Acacia, Sericomopsis,
sparse to open grassland in Fig. 1).
With increasing precipitation further south vegetation cover
becomes denser. With an erratic and unpredictable rainfall of
500e700 mm a year Acacia (A. tortilis, A. seyal, A. nilotica, A. senegal,
A. hockii, A. mellifera) dominated savanna grasslands have their
greatest development in a broad belt, which encircles Kenyas
 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
Fig. 1. Vegetation map of Kilimanjaro from Hemp (2006c), based on the evaluation of about 1400 vegetation plots following the method of Braun-Blanquet (1964) and a supervised
classification of Landsat ETM images taken on 29 January and 21 February 2000 (source: USGS/UNEP-GRID, Sioux Falls, SD, U.S.A.) using the software IDRISI 3.2 (Clark Labs,
Worcester, MA, U.S.A.). Digital elevation model by Christian Lambrechts and Janet Akinyi Onginjo, Nairobi, based on toposheets at scale1/1:50.000.
highlands, extending to Somalia and to north Tanzania (Tsavo,
Serengeti, Ngorongoro, Massai steppe; open grassland with sparse
shrubs, open open deciduous shrubland in Fig. 1). An increase in
rainfall favors wooded savanna grasslands dominated by e beside
Acacia - deciduous trees and bushes such as Commiphora, Termi-
nalia, Boscia, Lannea, Ozoroa, Sclerocarya and Grewia occurring in
Kenya, Uganda, Ruanda and Tanzania. The fruittree-like appearance
of the stunned woody species-mainly of belonging to Com-
bretaceae, Burseraceae and Anacardiaceae inspired the first bota-
nists to describe this vegetation formation as “Obstgartensteppe”
(fruitgardensteppe).
3.1.3. Miombo woodlands
With increasing tree height and cover these open vegetation
types change into woodlands, where trees reach a height of 15 m,
with the crowns just touching to form an open canopy, in contrast
to forest with a deeply closed canopy. Such Miombo or Brachyste-
gia- Julbernardia woodlands consists of deciduous trees such as
Brachystegia, Julbernardia and Isoberlinia, Afzelia, Terminalia, Lon-
chocarpus, Combretum and Markhamia (Burgess et al., 2004) and
have many species in common with savanna ecosystems but grades
into seasonal closed dry forest (Frost, 1996). Brachystegia-Julber-
nardia woodlands are one of the most extensive vegetation types in
Africa apart of the forests of Zaire (Lind and Morrison, 1974)
covering 2.7 million km2 and extending from Tanzania and south-
ern Democratic Republic of Congo (DR CONGO) in the north to the
northern provinces of South Africa, and across the continent from
Angola through Zambia to Malawi and Mozambique (Lind and
Morrison, 1974; Chidumayo, 1987). This vegetation formation
covers about two-thirds of Tanzania, whereas it is missing in Kenya
and Uganda. Miombo woodlands extend from sea level up to
1600 m in areas of 500e1200 mm annual rainfall in central and
southern Tanzania (Lind and Morrison, 1974) (deciduous woodland,
deciduous shrubland with sparse trees in Fig. 1). Burning and cut-
ting and conversion into agricultural fields and tobacco plantations
destroys rapidly increasing areas of Miombo woodlands.
303
3.1.4. Aquatic vegetation
Aquatic vegetation along shallow freshwater lake shores con-
sists of deep reed swamps, mainly of Papyrus (Cyperus papyrus),
other sedges and rushes such as Eleocharis, Typha domingensis, but
also grasses (Leersia, Oryza) and ferns (Thelypteris confluens).
Extensive Papyrus swamps can be seen at L. Victoria, L. Naivasha or
L. Jipe at the border between Tanzania and Kenya. The saline lakes
are fringed by marshes of Cyperus laevigatus. The floating and
submersed freshwater vegetation is characterized by Nymphaea,
Potamogeton, Ceratophyllum, Utricularia and Ottelia.
Surface of slowly floating rivers and canals, but also of sheltered
bays of lakes can be completely covered by the introduced water
lily (Eichhornia), giving problems in some areas, as well as Pistia
(Araceae) and water ferns such as Salvinia and Azolla.
3.1.5. Forests
Due to climatic restrictions and anthropogenic impact no more
than 5% of East Africa's surface is closed forest. These forests are
mainly confined to mountains. In the lowlands (below 800 m) only
few patches of forest remained, either occurring at the base of the
Eastern Arc Mts. and of the SW plateau of the Ethiopian highlands
or as remnants of the former widespread coastal forest strip
(evergreen lowland forest in Fig. 1).
The higher located montane forests of East Africa can be divided
into several elevational belts (nomenclature and delineation fol-
lows Hemp, 2006a). Apart from temperature, precipitation is the
main factor determining forest zonation (structure and species
composition). Submontane forests cover the foothills of the East
African mountains. Due to dense human settlements in such loca-
tions, only few patches in particular of moist evergreen forest are
left. At the Eastern Arc Mountains located in the low Tanzanian
coastal plains such forests start at 800e900 m, ranging to
1250e1400 m. Further inland, the submontane zone lies at about
1100e1700 mm. These submontane moist forest are botanically
very rich and consist of Allanblackia stuhlmannii, Cephalosphaera
usambarensis, Beilschmiedia kweo, Parinari excelsa and Newtonia
304 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
buchananii, receiving an over the year well-distributed rainfall of
more than 1100 mm.
In Uganda moist evergreen or semi-deciduous submontane
forests are now reduced by agricultural settlement to an archipel-
ago of small or medium forest reserves along the north-west shores
of Lake Victoria. At lower levels, from 1000 to 1200 m Cynometra
alexandri predominates, while from 1200 to 1700 m Parinari excelsa
is most frequent. Similar moist submontane forest types occur in
the isolated Kakamega forest in Western Province of Kenya, on the
SW escarpment of the NW Ethiopian highlands and in the lower
parts of the Harenna forest in the Bale Mts (Friis, 1986, 1992).
The montane forests in East Africa above 1600 m lie mainly on
the lower slopes of the major mountains, but also in the highlands
of Kigezi (Uganda) and Kenya, e.g. Cherengani Hills. On the wettest
mountains, in particular on the humid southern sides of the Kili-
manjaro and Kenya mountains, Eastern Arc, the dominant tree
species is camphor (Ocotea usambarensis; Lauraceae). Higher up
Podocarpus latifolius, Hagenia abyssinica (Rosaceae) and Prunus
africana (Rosaceae) form the tree canopy. On drier mountains, in
particular on the northern slopes the lower forests are character-
ized by Cassipourea malosana (Rhizophoraceae), higher up Junipe-
rus procera (Juniperaceae), Podocarpus latifolius and falcatus, and
Hagenia abyssinica are dominant tree species; such forest types
prevail in the highlands of Ethiopia and Somalia (Friis, 1992). A
typical component of most tall mountains in East Africa are dense
bamboo forests, which cover vast areas on the wet mountain slopes
above 2500 m, e.g. on the Kenya, Elgon, and Ruwenzori mountains,
the Virunga volcanoes and the Aberdare Range but not on Kili-
manjaro (submontane and montane forest in Fig. 1). Mount Kili-
manjaro in Tanzania has the widest gradient in elevation and
climate and can therefore serve as an example of forest zonation in
East Africa. The forest types of this mountain will be explained in
more detail below.
3.1.6. Alpine vegetation
The flora and vegetation of the tropical-alpine regions found
above the upper forest line is completely distinct from that of the
surrounding tropical lowlands (Hedberg, 1951; White, 1983). The
mountains on which this flora occurs form series of isolated ‘sky-
islands’ (Popp et al., 2008) or ‘archipelago-like regions’ (White,
1983; Harmsen et al., 1991). Alpine environments of East African
mountains are famous for their endemic and charismatic taxa, such
as the giant Lobelia, Dendrosenecio and shrubby Alchemilla with
very special growth forms such as giant rosettes.
The upper limit of the afroalpine belt is defined by the absence
of a vegetation cover, or the presence of a permanent snowline
(¼nival zone), which occurs at about 4500e5000 m. The lower
boundary of any alpine zone is commonly defined by the presence
of a forest line (Ko € rner and Paulsen, 2004; Ko € rner et al., 2011)
which is e.g. on Kilimanjaro at 4000 m. Between tree line and the
closed forest line normally lies a transitional vegetation zone of
several hundred meters in elevation. This subalpine zone between
the broad-leafed montane forest and the alpine Helichrysum scrub
vegetation (the treelineeecotone in the sense of Ko € rner, 2003)
corresponds closely to the “ericaceous belt” of (Hedberg, 1951) and
is a result of recurring fires or e on Mt. Elgon or in Ethiopia e of fire
and grazing (Hemp, 2005, 2006a).
The vegetation of the subalpine zone consists of ericaceous
shrublands, which represents different regeneration stages of burnt
forests. Due to the open canopy of the shrub layer (Erica arborea,
E. trimera, Protea caffra, Stoebe kilimandscharica, Anthospermum
usambarense and Adenocarpus mannii) many alpine species of the
alpine Helichrysum scrub occur. Between 3900 and 4000 m and
4600 m alpine dwarf scrub covers large areas, where Helichrysum
and Senecio are dominant and rich in species.
Altogether, East Africa sensu stricto (Kenya, Uganda, Tanzania) is
home to about 12.500 vascular plant species (FTEA), inhabiting a
huge variety of environments, ranging from semi-deserts to rain
forests, tropical coastal plains to alpine scrub vegetation.
3.2. Modern vegetation on Mount Kilimanjaro
Mount Kilimanjaro offers a great variety of vegetation types
ranging from dry succulent forests on the foothills at 800 m to
montane rainforests with a wealth of epiphytes and ferns to the
highest subalpine cloud forests of Africa at 4100 m as well as to the
alpine Helichrysum scrub (Fig. 2). Today's vegetation of Mt Kili-
manjaro harbours about vascular 3000 plant species (Hemp, 2006a,
2006b, 2006c).
3.2.1. Vegetation zonation
Based on significant floristic discontinuities in the composition
of the forests, the following forest zones can be distinguished on
the southern slope of Kilimanjaro (Hemp, 2006a): a colline
(¼lowland) zone (700e1100 m); submontane (1100e1700 m),
lower (1800e2100 m) and middle montane (2100e2800 m) forest
zones; an upper montane (2800e3200 m) forest zone; and a sub-
alpine (3200e4000 m) forest zone. Above the forestline the alpine
(4000e4600 m) and nival (>4600 m) zone follows. This terminol-
ogy is similar to that used by Lovett and Wasser (2008). On the basis
of fog water input and structure (e.g. richness of epiphytes) forests
of the middle, upper montane and subalpine zone can be defined as
‘cloud forests’. The elevational zones and the main vegetation types
viewed from the western side of the mountain (Fig. 2) demon-
strates the contrast between the dry northern and wet southern
slopes. The zonation is correlated with temperature and soil acid-
ity; and rainfall is of particular importance for zonation of epi-
phytes. Other key factors are humidity (influenced by stable cloud
condensation belts) and minimum temperature (in particular, the
occurrence of frost above 2700 m elevation; Hemp, 2006a).
3.2.2. Vegetation types
Owing to a large elevational range of over 5000 m and the
strong climatic contrast of the slopes, there is a large variety of
different vegetation types (Hedberg, 1951; Hemp, 2006a). Due to
drier climate conditions, the vegetation zones on the northern
slope of Kilimanjaro are shifted upslope and harbour a partly
different species composition.
Between 700 and 1100 m asl (1400e1600 m on the western and
northern slope) a dry and hot colline savanna zone stretches
around the mountain base. The mean annual temperature (MAT) is
between 20 and 24
C, and the mean annual precipitation (MAP)
reaches from 500 to about 1000 mm/a, hence, climatic conditions
are semiarid to subhumid. Today most of this area is used for crop
production (maize, bean and sunflower fields) or as pasture land.
On the north-western side of Kilimanjaro and around Lake Challa, a
deep crater lake in a secondary vent at the south-eastern foothills,
remnants of savanna woodlands with Acacia (Fabaceae), Grewia
(Tiliaceae/Malvaceae), Terminalia and Combretum (Combretaceae)
are still present. On the steep crater slopes of Lake Challa, dry for-
ests with the tree-succulents Euphorbia quinquecosta and
E. candelabrum (Euphorbiaceae) as well as broadleaved deciduous
species such as Commiphora baluensis (Burseraceae) and Hap-
locoelum foliosum (Sapindaceae) can be found. Such steep volcanic
cones together with the narrow bands of riparian forests with
species such as Ficus sycomorus (Moraceae) and Lecaniodiscus
fraxinifolius (Sapindaceae) represent the only patches of natural
vegetation in most of the former savanna area.
Due to the favourable climate (MAT: 16e20
C, MAP:
1000e2000 mm), today the submontane zone of the southern and
 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386 305

Fig. 2. Land-cover map of East Africa modified from Bartholome and Belward (2005); flora regions (T1-8; K1-7; U1e4) covered in this atlas are projected according to FTEA
(1955e2012). The landcover map is a product of the Global Land Cover 2000 project (\\BECKINSALE\GLC2000\Africa\V5\Grid\africa_v5), coordinated by the Global Vegetation
Monitoring Unit of the European Commission Joint Research Center. The land-cover data are derived in 2000 from the VEGETATION instruments onboard the SPOT4 satellite, the
ATSR-2 and the JERS-1.

eastern slope is the main region of agri- and horticulture with ba-
nana and coffee plantations that extend from 1000 up to 1800 m.
This zone has been inhabited by the Chagga, a Bantu tribe, for
several centuries. The Chagga cultivate their land using an agro-
forestry system (“Chagga homegardens”). Remnants of the former
forests of this zone have survived only in the deepest valleys and
gorges. These forests, although of small extent, are of great
biogeographical and palaeobotanical importance. They differ
entirely in species composition and structure from the forests of
higher elevations, resembling instead the diverse wet montane
forest of the Pare and Usambara mountains. The forest vegetation is
highly divers and includes species such as Entandrophragma
excelsum (Meliaceae, up to over 80 m), Heinsenia diervilleoides,
Hallea rubrostipulata (all Rubiaceae), Newtonia buchananii (Mim-
osaceae), Leptonychia usambarensis (Sterculiaceae), Strombosia
scheffleri (Loganiaceae), Dasylepis integra (Flacourtiaceae), Garcinia
tanzaniensis (Clusiaceae) and Polyscias albersiana (Araliaceae). In
the western parts of the mountain, the comparatively dry sub-
montane forest below 1600 m is dominated by Olea europaea ssp.
africana (Oleaceae), Croton megalocarpus (Euphorbiaceae), Calo-
dendrum capense (Rutaceae) and Diospyros abyssinica (Ebenaceae).
The montane forests of Mt Kilimanjaro border the cultivated
zone on the southern and southeastern slopes between 1700 and
1800 m and surround the whole mountain. Within the montane
forest zones three sub-zones can be distinguished: lower, middle
and upper montane forest.
The lower montane zone on the southern slope ranges from
1700 to 2100 m asl (MAT: 14.4e16
C, MAP: 2000e2700 mm).
Above 1700 m Ocotea usambarensis (Lauraceae) is associated with
Agarista salicifolia (Ericaceae), Syzygium guineense (Myrtaceae),
Macaranga capensis var. kilimandscharica (Euphorbiaceae), and
Polyscias fulva (Araliaceae).
The middle montane forest zone (2100e2800 m asl) receives
the highest amount of rainfall on the whole mountain (MAP:
1910e3050 mm) and has a MAT of 10.4e14.4
C. These favourable
climate conditions give rise to a high richness in ferns and epi-
phytes. The frost susceptible tree fern Cyathea manniana is very
abundant below 2700 m asl which is the elevation at which frost
starts to occur occasionally during the night. In the lower part of
this belt the camphor-tree Ocotea usambarensis dominates together
with Ilex mitis (Aquifoliaceae) and Xymalos monospora (Mon-
imiaceae), in the upper part Ocotea is increasingly associated with
Podocarpus latifolius (Podocarpaceae). In the drier southwestern
part of the camphor belt, Ocotea forests with the tree Faurea
wentzeliana (Proteaceae) exist at this altitude. In gorges and along
streams Cornus volkensii (Cornaceae) is an important constituent of
the tree layer.
Above 2800 m asl Podocarpus latifolius (Podocarpaceae) is the
characteristic dominant tree of the upper montane forest zone
together with Hagenia abyssinica (Rosaceae) and Prunus africana
(Rosaceae). Monodominant stands of Erica excelsa (Ericaceae) play
also an important role in this zone, replacing Podocarpus and
Hagenia forests after fire, forming the actual upper closed forest line
on the southern slope of Kilimanjaro at 3250 m (Hemp and Beck,
2001). However, small remnants and burnt forests indicate that
the upper closed forest line reached up to 3850 m recently and
remnants of subalpine Erica trimera forests mark the former and
potential upper closed forest line at above 4000 m (Hemp, 2005),
representing today the highest elevation forests in Africa.
Due to the drier climate, in particular on the lower slopes,
different forest types occur on the northern side of the mountain. In
the submontane zone (1600e2000 m asl) a similar Croton-
306 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386

Calodendrum forest grows as on the western slope. In the lower
montane forest zone (2000e2500 m asl) Cassipourea malosana
(Rhizophoraceae), Teclea simplicifolia, Fagaropsis angolensis (Ruta-
ceae) and Olea capensis are abundant. Above 2500 m up to 3100 m
asl, a Juniperus-Podocarpus-forest (Juniperus procera, Cupressaceae,
Podocarpus latifolius, P. falcatus, Nuxia congesta, Hagenia abyssinica)
replaces the lower montane forest. After fire monospecific stands of
Erica excels substitute Juniperus-Podocarpus-forests. With
increasing elevation climatic conditions (in particular rainfall)
become similar to the conditions on the southern slope; therefor
vegetation above 3100 m asl, the subalpine Erica-shrubland as well
as the alpine and nival zones harbour a similar flora as found on the
southern slope.
In the subalpine zone e the treeline ecotone in the sense of
Ko€rner 2003 - between 3200 and 4000 m the forests of Erica excelsa
are gradually replaced by Erica-shrub with the dominant species
Erica arborea and Erica trimera, Protea caffra and Euryops dacry-
dioides. In the southeastern parts, moorland vegetation, formed by
tussock grass and characterized by giant lobelias, fringes the forest.
In contrast to the dry moorlands Carex bogs e an important habitat
of Dendrosenecio - occur on wet sites along streams or water
sources. MAT is between 4.2 and 8.7
C. This belt receives less
rainfall (MAP: 700e1500 mm) which facilitates the occurrence of
fires.
The alpine zone begins at an altitude of about 4000 m and here,
the Erica heathlands grade into a Helichrysum cushion vegetation
that extends up to 4600 m. Due to the rather hostile climate con-
ditions (MAT: 4.2 to 0.7
C, MAP: 500e700 mm) these altitudes are
poor in vegetation. A dwarf scrub with cushions of Helichrysum
newii and H. citrispinum, Senecio meyeri-johannis (all Asteraceae)
and Pentaschistis borussica (Poaceae) covers large areas.
In the nival zone MAT is below 0
C and annual precipitation is
low (MAP: <500 mm). The boulder slopes are only sparsely vege-
tated by lichens and mosses. The top of Kibo (5895 m asl) is still
partly covered with glaciers.
Most of the former savanna vegetation on the foothills of Kili-
manjaro is converted into agricultural fields with maize, sunflowers
and beans and weeds such as Malvastrum coromandelianum and
Sida species. Higher up in the submontane zone traditional mixed
coffee and banana plantations cover most of the area below the
forest belt at 1700e1800 m. These Chagga homegardens represent
a special type of agro-forestry with a multi-layered vegetation
structure similar to that of a tropical montane forest, with forest
trees such as Albizia schimperiana (Mimosaceae), Bridelia micrantha
(Euphorbiaceae), as well as Mango and Papaya trees, shrubs, lianas,
epiphytes and herbs.
4. Material and method
4.2. Photo documentation of pollen and spores
Images were obtained using Leica Microscope and the Leica DM
6000 Band camera with the accompanying image analysis Leica
QCapture software and a Rollei Compactline 110 camera. A scale bar
of 10 mm was added to at least one picture of each pollen and spore
type.
Photos of all identified pollen and spores as well as the original
slides are available on request at the Department of Palynology and
Climate Dynamics (Albrecht-von-Haller-Institute for Plant Sci-
ences, University of Go €ttingen, Germany). Pollen grain measure-
ments represent approximate values which were obtained by
measuring ~30 grains per pollen type. Due to intraspecific variation
in pollen grain size in some taxa (e.g. Joly et al., 2007; Lau and
Stephensen, 1993), Sosnoskie et al., 2009) not all grain size values
can be used as reliable discriminating character. However, our
experience has shown that for most pollen types with large dif-
ferences in size (5e10 mm) the measurement values definitely
help the identification.
4.3. Pollen and spore identification and nomenclature
The nomenclature of the herbarium specimens as well as the
field collections follows the Flora of Tropical East Africa (FTEA,
1955e2012; Lovett and Wasser, 1993; Beentje et al., 1994; Hemp,
2001a, 2006a). The pollen and spore terminology follows Punt
et al. (2007). According to Punt et al. (2007), the term ‘circular’
describes the pollen grain shape in polar view, whereas the term
‘sphoidal’ describes the pollen grain shape in equatorial view; this
concept is also applied in this atlas. If possible, pollen and spore
morphology was verified with relevant literature (Erdtman, 1957;
Maley, 1970; Ybert, 1979; Iwatsuki, 1990; Tryon et al., 1990;
Roubik and Moreno, 1991; Tryon and Lugardon, 1991; Verdcourt,
zine, 2001, Verdcourt and Beentje, 2001a, 2001b;
1999a, 1999b; Le
Verdcourt, 2002, 2003, 2005a, 2005b, 2006; Roux et al., 2007;
Vincens et al., 2007; Gosling et al., 2013, and other literature as
mentioned in the entries).
Pollen and spore types are ordered taxonomically by family
following FTEA. The correct taxonomy of species was verified with
the online African Plant Database (APD, Version 3.4.0). Changes in
taxonomic affiliation on family level are indicated behind the
family or species name. All described pollen (Table 1) and spore
types (Table 2) indicate the plate number on which the corre-
sponding images are displayed.
Table 1
List of described spore types, taxonomic affiliation and corresponding
plates with the spore illustrations.

Family Species Plate

4.1. Pollen collection and sample preparation Actiniopteridaceae Actiniopteris dimorpha I, 1e2
Adiantaceae/ Adiantum capillus-veneris I, 3e5
All pollen and spores were obtained from the reference collec- Pteridaceae Adiantum incisum I, 6e7
tion of ca. 2000 specimens housed within the Department of Adiantum poiretii var. poiretii I, 8e10
Coniogramme africana I, 11e12
Palynology and Climate Dynamics, Go €ttingen of University, which
Pellaea dura var. dura I, 13e15
has been collected and curated by L. Schüler and A. Hemp. Refer- Pityrogramma aurantiaca I, 16e18
ence material was sourced from the East Africa herbarium of A. Aspleniaceae Asplenium adiantum-nigrum II, 1e2
Hemp at the Department of Plant Systematics at the University of Cyatheaceae Cyathea humilis II, 3e5
Bayreuth (herbarium code: UBT) and field collections in the Kili- Cyathea manniana II, 6e7
Dryopteridaceae Arachniodes webbiana subsp. II, 8e10
manjaro area between 2009 and 2013. The pollen grains were foliosa
extracted from dried and fresh plant material by sieving the sam- Didymochlaena truncatula II, 11e12
ples. Afterwards, acetolysis was carried out on the samples to Gleicheniaceae Dicranopteris linearis II, 13
dissolve cellulose material and to stain the pollen residues (Moore Grammitidaceae/ Xiphopteris flabelliformis II, 14e16
Polypodiaceae
et al., 1991). The pollen residue was kept in distilled water until
mounted into glycerol jelly for the preparation of pollen slides.
 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386 307

Table 1 (continued ) Table 2 (continued )

Family Species Plate Family Species Plate

Hymenophyllaceae Hymenophyllum capillare II, 17e18 Helichrysum forskahlii var. VI, 31e33
Trichomanes melanotrichum II, 19e20 forskahlii
Lomariopsidaceae/ Elaphoglossum acrostichoides II, 21e23 Helichrysum schimperi VI, 34e36
Dryopteridaceae Hirpicium diffusum VI, 37e38
Lycopodiaceae Huperzia dacrydioides II, 24e25 Lactuca capensis VII, 1e2
Huperzia ophioglossoides II, 26e27 Pluchea nitens VII, 3e5
Huperzia saururus II, 28e29 Schkuhria pinnata VII, 6e8
Lycopodium clavatum II, 30e32 Senecio cyaneus VII, 9e12
Marattiaceae Marattia fraxinea II, 33e34 Senecio johnstonii VII, 13e16
Ophioglossaceae Ophioglossum vulgatum subsp. II, 35e37 Siegesbeckia abyssinica VII, 17e18
kilimandscharicum Sphaeranthus bullatus VII, 19e20
Polypodiaceae Loxogramme lanceolata II, 38e39; Spilanthes mauritiana VII, 21e23
III, 1 Stoebe kilimandscharica VII, 24e26
Lepisorus excavatus III, 2e4 Tagetes minuta VII, 27e29
Pteridaceae Pteris dentata III, 5e8 Tolpis capensis VII, 30e32
Selaginellaceae Selaginella goudotiana var. III, 9e11 Tridax procumbens VIII, 1e3
abyssinica Vernonia brachycalyx VIII, 4e6
Thelypteridaceae Amauropelta bergiana III, 12e14 Vernonia colorata VIII, 7e8
Christella dentata III, 15e17 Vernonia lasiopus VIII, 9e10
Woodsiaceae/ Cystopteris fragilis ssp. diaphana III, 18e20 Vernonia syringifolia VIII, 11e12
Cystopteridaceae Balanitaceae Balanites aegyptiaca VIII, 13e15
(Zygophyllaceae) Balanites maughamii ssp. acuta VIII, 16e17
Balsaminaceae Impatiens pseudoviola VIII, 18e19
Impatiens walleriana VIII, 20e21
Table 2 Basellaceae Basella alba VIII, 22e24
List of described pollen types, taxonomic affiliation and corresponding plates Begoniaceae Begonia johnstonii VIII, 25e26
with the pollen illustrations. Bignoniaceae Jacaranda mimosifolia VIII, 27e29
Family Species Plate Markhamia lutea VIII, 30e31 &
IX, 1
Acanthaceae Asystasia gangetica IV, 1e2 Tecoma stans IX, 2e3
Blepharis maderaspatensis IV, 3e5 Bombacaceae Adansonia digitata IX, 4e5
Isoglossa laxa IV, 6e9 (Malvaceae)
Justicia asystasioides IV, 10e12 Boraginaceae Cordia monoica IX, 6e7
Justicia diclipteroides IV, 13e17 Ehretia bakeri IX, 8e10
Thunbergia alata IV, 18e20 Heliotropium rariflorum ssp. IX, 11e12
Amaranthaceae Achyranthes aspera IV, 21e23 hereroense
Amaranthus hybridus IV, 24e26 Lithospermum afromontanum IX, 13e14
Celosia schweinfurthiana IV, 27e28 Burseraceae Boswellia neglecta IX, 15e17
Anacardiaceae Lannea triphylla IV, 29e31 Caesalpiniaceae Caesalpinia decapetala IX, 18e20
Ozoroa insignis IV, 32e35 Cassia spectabilis IX, 21e24
Rhus vulgaris IV, 36e39 Delonix regia IX, 25e27
Anthericaceae Chlorophytum rhizopendulum IV, 40e41 Pterolobium stellatum IX, 28e29
Chlorophytum viridescens IV, 42e44 Campanulaceae Wahlenbergia abyssinica IX, 30e31 & X,
ApocyNaceae Carissa edulis IV, 45e47 1
Rauvolfia caffra IV, 48e49 Capparaceae Cleome hirta X, 2e3
Saba comorensis IV, 50e52 Maerua grantii X, 4e6
Tabernaemontana stapfiana IV, 53 & V, 1 Caryophyllaceae Cerastium indicum X, 7e8
e3 Uebelinia rotundifolia X, 9e10
Aquifoliaceae Ilex mitis var. mitis V, 4e6 Celastraceae Hippocratea africana X, 11e13
Araliaceae Cussonia arborea V, 7e8 Maytenus acuminata X, 14e16
Polyscias fulva V, 9e11 Chenopodiaceae Chenopodium ambrosioides X, 17e18
Schefflera myriantha V, 12e15 Combretaceae Combretum apiculatum X, 19e21
Schefflera volkensii V, 16e18 Combretum psidioides subsp. X, 22e24
Asparagaceae Asparagus racemosus V, 19e20 psidioides
Asteraceae Artemisia afra V, 21e22 Commelinaceae Aneilema aequinoctiale X, 25e26
Bidens pilosa V, 23e25 Aneilema johnstonii X, 27e28
Bothriocline amplifolia V, 26e28 Commelina foliacea X, 29e30
Bothriocline longipes V, 29e31 Connaraceae Rourea thomsonii X, 31e33
Carduus keniensis V, 32e33 Convolvulaceae Astripomoea hyoscyamoides X, 34e35
Centaurea praecox V, 34e36 subsp. hyoscyamoides
Cineraria deltoidea V, 37e39 Convolvulus kilimandschari X, 36e38
Conyza newii VI, 1e2 & V, Crassulaceae Crassula alsinoides X, 39e42
40 Cucurbitaceae Diplocyclos schliebenii XI, 1e3
Conyza ruwenzoriensis VI, 3e6 Luffa cylindrica XI, 4e6
Conyza vernonioides VI, 7e8 Momordica boivinii XI, 7e9
Crassocephalum bojeri VI, 9e11 Momordica foetida XI, 10e11
Crepis oliveriana VI, 12e13 Oreosyce africana XI, 12 & XII, 1
Dicoma tomentosa VI, 14e16 Cyperaceae Carex conferta XII, 2e3
Dicrocephala integrifolia VI, 17e18 Carex vallis-rosetto XII, 4e6
Erlangea calycina VI, 19e20 Cyperus amauropus XII, 7e9
Gerbera viridifolia VI, 21e23 Cyperus rigidifolius XII, 10e11
Haplocarpha rueppellii VI, 24e25 Dracaenaceae Dracaena afromontana XII, 12e13
Helichrysum argyranthum VI, 26e28 Dracaena fragrans XII, 14e15
Helichrysum citrispinum VI, 29e30 Ebenaceae Euclea divinorum XII, 16e18
(continued on next page) (continued on next page)
308 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386

Table 2 (continued ) Table 2 (continued )

Family Species Plate Family Species Plate

Ericaceae Agauria salicifolia XII, 19e21 Piperaceae Peperomia blanda XVII, 28e29
Erica arborea XII, 22e24 Piper capense var. capense XVII, 30e31
Euphorbiaceae Clutia abyssinica var. abyssinica XII, 25 & XIII, 1 Plantaginaceae Plantago palmata XVII, 32e34
e2 Plumbaginaceae Plumbago dawei XVII, 35e37
Croton megalocarpus XIII, 3e4 Poaceae Cynodon dactylon XVII, 38e39
Euphorbia engleri XIII, 5 Dactyloctenium aegyptium XVIII, 1e2
Euphorbia schimperiana var. XIII, 6e8 Diheteropogon amplectens XVIII, 3e4
schimperiana Eleusine jaegeri XVIII, 5e6
Macaranga kilimandscharica XIII, 9e10 Elionurus muticus XVIII, 7e8
Phyllanthus boehmii var. boehmii XIII, 11e12 Harpachne schimperi XVIII, 9e10
Fabaceae Aeschynomene schimperi XIII, 13e15 Hyperthelia dissoluta XVIII, 11e13
Calpurnia aurea XIII, 16e17 Isachne mauritiana XVIII, 14e15
Crotalaria axillaris XIII, 18e20 Podocarpaceae Podocarpus falcatus XVIII, 16e18
Crotalaria goodiformis XIII, 21e23 Podocarpus latifolius XVIII, 19e21
Crotalaria keniensis XIII, 24e26 Polygonaceae Polygonum salicifolium XVIII, 22e23
Dalbergia lactea XIII, 27e28 Rumex bequaertii XVIII, 24e26
Desmodium repandum XIII, 29e31 Proteaceae Grevillea robusta XVIII, 27e28
Eriosema montanum var. XII, 32e34 Protea kilimandscharica XVIII, 29e30
montanum Ranunculaceae Clematis brachiata XVIII, 31e32 &
Erythrina burtii XIII, 35e37 XIX, 1
Glycine wightii agg. XIV, 1e3 Ranunculus oreophytus XIX, 2e4
Indigofera atriceps subsp. XIV, 4e6 Thalictrum rhynchocarpum XIX, 5e6
kaessneri Resedaceae Caylusea abyssinica XIX, 7e9
Indigofera swaziensis subsp. XIV, 7e8 Rhamnaceae Scutia myrtina XIX, 10e12
swaziensis Rosaceae Alchemilla volkensii XIX, 13e16
Flacourtiaceae Dasylepis integra XIV, 9e12 Hagenia abyssinica XIX, 17e18
Dovyalis abyssinica XIV, 13e15 Rubiaceae Chassalia kenyensis XIX, 19e21
Rawsonia lucida XIV, 16e18 Chassalia parviflora XIX, 22e25
Hamamelidaceae Trichocladus ellipticus XIV, 19e20 Galiniera saxifraga XIX, 26e28
Hypericaceae (partly Garcinia volkensii XIV, 21e23 Kohautia coccinea XIX, 29e31
Clusiaceae) Hypericum revolutum ssp. XIV, 24e26 Lasianthus kilimandscharicus XIX, 32e34
keniense Mussaenda microdonta XIX, 35 & XX,
Icacinaceae Apodytes dimidiata XIV, 27e28 1
Iridaceae Gladiolus watsonioides XIV, 29e30 Pavetta abyssinica var. abyssinica XX, 2e4
Lamiaceae Clerodendron johnstonii XIV, 31e33 Pentas lanceolata ssp. lanceolata XX, 5e9
Leonotis mollissima XIV, 34e36 var. lanceolata
Leucas volkensii XIV, 37e40 Pentodon pentandrus var. XX, 10e12
Plectranthus alboviolaceus XIV, 41e42 pentandrus
Plectranthus laxiflorus XIV, 43e44 & Psychotria capensis ssp. riparia XX, 13e14
XV, 1 Psychotria cyathicalyx XX, 15e16
Plectranthus sylvestris XV, 2e4 Psychotria fractinervata XX, 17e18
Lauraceae Ocotea usambarensis XV, 5e6 Psychotria petiginosa XX, 19e20
Linaceae Linum volkensii XV, 7e8 Richardia scabra XX, 21e22
Lobeliaceae Cyphia glandulifera XV, 9e11 Rutaceae Calodendrum capense XX, 23e25
(Campanulaceae) Lobelia deckenii XV, 12e14 Clausena anisata XX, 26e28
Lobelia holstii XV, 15e17 Toddalia asiatica XX, 29e31
Loganiaceae (partly Nuxia congesta XV, 18e21 Santalaceae Osyris lanceolata XX, 32e35
Stilbaceae) Strychnos scheffleri XV, 22e23 Sapindaceae Allophylus ferrugineus XX, 36e37
Loranthaceae Agelanthus elegantulus XV, 24e25 Allophylus rubifolius var. XX, 38e39
Englerina woodfordioides XV, 26e27 rubifolius
Plicosepalus curviflorus XV, 28e30 Filicium decipiens XXI, 1e3
Tapinanthus XV, 31e32 Paullinia pinnata XXI, 4e5
Malvaceae Abutilon longicuspe var. XV, 33e35 Sapotaceae Aningeria adolfiefriedericii agg. XXI, 6e8
longicuspe Solanaceae Solanum nigrum XXI, 9e11
Hibiscus meyeri XVI, 1e2 Solanum seaforthianum XXI, 12e14
Hibiscus adoensis XVI, 3e4 Sterculiaceae Dombeya burgessiae XXI, 15e16
Pavonia elegans XVI, 5e6 (Malvaceae) Leptonychia usambarensis XXI, 17e19
Sida ovata XVI, 7e9 Thymeliaceae Gnidia apiculata XXI, 20e21
Meliaceae Lepidotrichilia volkensii XVI, 10e12 Gnidia glauca XXI, 22e23
Trichilia emetica XVI, 13e15 Gnidia subcordata XXI, 24e25
Turraea holstii XVII, 1e2 Tiliaceae (Malvaceae) Grewia bicolor XXI, 26e28
Menispermaceae Stephania abyssinica XVII, 3e5 Grewia microcarpa XXI, 29e31
Mimosaceae Acacia polyacantha var. XVII, 6e7 Triumfetta brachyceras XXI, 32e34
camplyacantha Triumfetta flavescens XXI, 35e37
Albizia schimperiana var. XVII, 8e9 Ulmaceae Celtis gomphophylla XXII, 1& XXI,
amaniensis (Cannabaceae) 38e40
Mimosa invisa XVII, 10e12 Trema guineensis XXII, 2e3
Monimiaceae Xymalos monospora XVII, 13e14 Urticaceae Elatostemma paivaeanum XXII, 4e5
Myrsinaceae Rapanea melanophloeos XVII, 15e17 Pilea johnstonii XXII, 6e7
(Primulaceae) Urera hypselodendrum XXII, 8e9
Myrtaceae Syzygium guineense ssp. XVII, 18e19 Valerianaceae Valeriana volkensii XXII, 10e12
afromontanum (Caprifoliaceae)
Oleaceae Jasminum abyssinicum XVII, 20e22 Vitaceae Cissus oliveri XXII, 13e15
Olea capensis ssp. hochstetteri XVII, 23e24 Cyphostemma kilimandscharicum XXII, 16e19
Oxalidaceae Oxalis latifolia XVII, 25e27
 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386 309

The data on distribution (Fig. 1) follows the information given in
the FTEA (1955e2012). Yet unpublished occurences in the Kili-
manjaro area (T2) (Hemp, unpublished data) were added accord-
ingly. The distribution and habitat description at Kilimanjaro is
derived from detailed vegetation surveys by Hemp during the past
20 years (Hemp, 2001a, 2001b, 2002, 2005a; 2005b; 2006a, 2006b;
2006c, 2008). The distribution categories for Kilimanjaro
(extremely rare, very rare, rare, scattered, widespread) are based on
the occurrence in the number of 1 km2 UTM grids (1e10, 11e25,
26e50, 51e150, >150).
4.4. Pollen key
The pollen key (Table 3) includes all pollen types described in
this atlas and fascilitate the correct indentification of pollen grain
based on distinct morphological characteristics. An identication key
for spores was not compiled due to the relatively limited number of
fern taxa included in the atlas.

Table 3
Pollen key to the pollen types described in this atlas.

1) Grain Arrangement
➢ Monade /5
➢ Monade, vesiculate /6
➢ Tetrade /3
➢ Polyade (8 or 16 grains) /2
2) Polyade
➢ Polyad (Ø ~82 mm) with 16 rectangular grains, 6e8 porous areas or faint colpi, finely scabrate Albizia schimperiana var. a.
➢ Polyads (Ø ~38 mm) of (8 or) 16 grains, single grains (~11 mm) pyramida, psilate Acacia polyacantha var. c.
3) Tetrade
➢ 3-colporate /4
➢ 4-porate, pori with thin annulus, scabrate, tetrade Ø ~25 mm Mimosa invisa
4) 3-colporate tetrade
➢ Psilate to finely granulate, tetrad Ø ~27 mm, narrow colpi with distinct costae, lalongate pori Agauria salicifolia
➢ Rugulate, tetrad Ø 30 mm, narrow colpi with distinct costae but often wider in pore area, lalongate pori Erica arborea
5) Monade
➢ Inaperturate /7
➢ 1 aperture /8
➢ 2 apertures /11
➢ 3 apertures /12
➢ 4 apertures /15
➢ 5 apertures /16
➢ 6 apertures /17
➢ Poly/peri/stephanoperaturate /18
➢ Spiraperturate /19
6) Vesiculate monads
➢ Corpus (27e33 x 23e29 mm) psilate to finely scabrate, the sacci reticulate, muri high Podocarpus falcatus
➢ Corpus (22x44  32 mm) coarsely scabrate to granulate, the sacci (incompletely) reticulate, muri often with open ends Podocarpus latifolius
7) Inaperturate monads
➢ Clavate, (sub)prolate ~27  38 mm, circular to elliptic Xymalos monospora
➢ Microechinate, spheroidal and circular but often deformed ~ 32 mm Ocotea usambarensis
➢ Reticulate, ‘crotonoid’, circular, spheroidal ~60 mm Croton megalocarpus
➢ Verrucate, circular, spheroidal ~11 mm Peperomia blanda
8) Monoaperturate Monads
➢ Colpate/sulcate /9
➢ Porate /10
9) Monocolpate/sulcate
➢ Bacculate
B Colpus very broad and long (5/6), loosely bacculate with relative large bacculae, grain ± circular, ~55 mm Aneilema aequinoctiale
B Colpus narrow to broad about 2/3 long, densely bacculate with small bacculae, grain subprolate ~ 50  32 mm Aneilema johnstonii
➢ Microechinate, subprolate, broad colpus with granulate margins Gladiolus watsonioides
➢ Psilate/scabrate
B Subprolate ~ 16  28 mm, colpus narrow with grainy margins Chlorophytum rhizopendulum
B Subprolate~ 27  36 mm, colpus wider in the center Chlorophytum viridescens
B Prolate ~20  32 mm, ± circular, colpus (3/4 to 4/5) mostly broad Asparagus racemosus
➢ Reticulate/microreticulate
B Colpus 5/5 wide, sub-prolate ~44  69 mm to spheroidal, ± circular ~57 mm, finely reticulate Dracaena afromontana
B Colpus 5/5, wide (<than in D. afrom.), (sub)prolate ~44  66 mm, ± circular, reticulate (-pilate) Dracaena fragans
B Colpus (3/4 to 4/5) mostly broad, prolate ~20  32 mm, ± circular, microreticulate Asparagus racemosus
➢ Microverrucate
B Circular grain (~10e12 mm), densly verrcuate Piper capense var. c.
B Broad colpus with fissured margins, subprolate (~24 mm) Commelina foliacea
10) Monoporate
➢ Psilate/scabrate, grains ± circular and spheroidal
B Pore Ø ~2 mm, annulus width ~3.5 mm protruding distinctly, grain ~33 mm Diheteropogon amplectens
➢ Scabrate/granulate/microechinate, grains ± circular and spheroidal
B Circular pore with annulus and operculum
▪ Pore Ø ~3 mm, annulus ~3 mm wide slightly protruding, coarsely scabrate, grain ~37 mm Hyperthelia dissoluta
▪ Pore and annulus (costa & margo) not protruding, pore Ø~2.4e3.3 mm, annulus ~1.6e1.7 mm wide, compactly insular exine pattern, Cynodon dactylon
grain ~30e33 mm
▪ Annulus with thick costa and thin margo, pore Ø ~2.5 mm in level with ectexine or slightly sunken, sparsely insular exine pattern, Eleusine jaegeri
grain~25 mm
(continued on next page)
310 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386

Table 3 (continued )

B Circular pore circular with annulus, without operculum

▪ Annulus protruding distinctly, pore Ø ~3 mm, annulus ~3 mm wide, grain~40 mm Elionurus muticus
▪ Annulus protruding, pore circular Ø ~2 mm, annulus ~ 1.5 mm wide, grain ~31 mm Isachne mauritiana
▪ Pore slightly sunken Ø 4 mm, annulus width ~2 mm, microechinate, grain ~31 mm Dactyloctenium aegyptium
▪ Pore Ø ~ 3.5 mm, faint annulus ~3 mm wide, coarsely scabrate Harpachne schimperi
11) Diaperturate
➢ 2-porate
B Loosely microechinate, circular pori with annulus, grain circular ~ 14 mm Pilea johnstonii
B (Micro)reticulate, pore with marginal girdle, very thick exine Isoglossa laxa
B Scabrate, circular pore with slightly protruding annulus, grains circular ~ 11 mm Elatostemma paivaeanum
B Bacculate/granulate, circular-elliptic pore with thick annulus, grain circular, spheroidal, ~25 mm Celtis gomphophylla
➢ 2-colporate
B Reticulate, Pore elliptic broad with well-defined costa, colpi long and narrow,grain rectangular/prolate ~38  23 mm, crassinexious Justicia diclipteroides
12) Triaperturate
➢ 3-porate /13
➢ 3-colpate/colporate /14
13) 3-porate
➢ Echinate
B pori lalongate with thin annulus, columellae distinct, spines short, pointed with a small basal cushion Abutilon longicuspe var. l.
B pori circular with annulus, spines short pointy, base distinctly broader Dombeya burgessiae
B grains circular ~83 mm, large ± circular pori, distinctly foveolate, loosly echinate Diplocyclos schliebenii
B microechinate, circular pori, grains circular ~26 mm Wahlenbergia abyssinica
➢ Psilate
B circular pori with protruding annulus, circular, spheroidal, ~24 mm Trema guineensis
B large pori Ø~ 15 mm with thin annulus and small vestibulum, circular, spheroidal ~75e91 mm Oreosyce africana
B pori circular Ø 1e1.5 mm with thin annulus which is not protruding, spheroidal, circular; ~13 mm Urera hypselodendrum
➢ Reticulate, reticuloid
B Grain ± circular and spheroidal/subprolate
▪ pori ± circular, small annulus, coarse reticulum, circular, spheroidal ~12 mm Stephania abyssinica
▪ pori circular, annulus slightly protruding, microreticulate, circular, spheroidal ~24 mm Calodendrum capense
▪ large pori rectangular ~3  7 mm, grain circular to subtriangular ~35 mm, subprolate Lasianthus kilimandscharicus
B Grain ± triangular
▪ circular pori large Ø ~8 mm, reticulate with large-sized lumina and very broad muri, grain triangular, subprolate to rhombic tall Glycine wightii
~35  40 mm
▪ pori elliptic tall (annulus?), reticulate with large lumina, grain triangular (convex) ~31 mm, (sub)oblate Leptonychia usambarensis
▪ 3 (4) porate, large elliptic pori, finely reticulate, grain triangular ~ 44 mm, (sub)oblate Paullinia pinnata
▪ pori large Ø 11 mm, surface undulating/reticuliod pattern, triangular ~57 mm, oblate Grevillea robusta
➢ Striato-reticulate, grains triangular ~21 mm, oblate, pori circular with distinct margins Allophylus rubifolius var. r.
➢ Scabrate, grains triangular ~34 mm, oblate, exine thickened around pori, intine around pori bent inwards Allophylus ferrugineus
➢ Verrucate/bacculate, circular grain ~63 mm, ± circular pori with annulus Adansonia digitate
➢ Bacculate/granulate, circular grain, circular-elliptic pore with annulus, circular, spheroidal, ~25 mm Celtis gomphophylla
14) 3-colpate/col(porid)ate/colporate
➢ Aerolate
B 3-colporate, aerolate/verrucate, triangular in polar view Dicoma tomentosa
➢ Bacculate
B 3- colporate, pori with operculum, pollen grain circular Filicium decipiens
➢ Clavate
B 3-colpate, circular grain ~35 mm, deeply recesses colpi Linum volkensii
B 3-colpor(id)ate, clavae heteromorphic, grain triangular ~ 14 mm Ilex mitis var. m.
➢ Echinate/microechinate
B Microechinate (spines <1 mm)
▪ 3-colpate
B Subprolate, trilobate, colpi (5/6) with obtuse ends Clematis brachiata
B (Sub)oblate, distinctly three-armed, syncolpate Agelanthus elegantulus
B Circular (~60 mm), subprolate, straight colpi ¾ in length with irregular margins and obtuse ends Clerodendron johnstonii
▪ 3-colpor(id)ate
B Pore elliptic tall or lolongate
▪ colpi 2/3 and narrow, pori elliptic, grain spheroidal, circular ~46 mm Cordia monoica
▪ colpi 5/6 and narrow, pori elliptic, distinctly double-columellate, grain subprolate-spheroidal, circular ~25 mm Artemisia afra
▪ recessed colpi long and open, large pori lolongate, grain ~35 mm Senecio johnstonii
B Pore elliptic broad/lalongate
▪ long (5/6) and broad colpi, pore elliptic broad tapering off lalongate, exine thinner towards apertures, grain 32  29 mm Centaurea praecox
B Pore circular or inconspicuous
▪ colpi (5/6) wide at the equator, acute ends, thin margo, grain ~40 mm Valeriana volkensii
▪ syncolpate, narrow colpi with margo, circular pori, triangular concave,longest axis ~33 mm, oblate Plicosepalus curviflorus
B Echinate (spines >1 mm), 3-colpor(id)ate
▪ Grains ≤ 34 mm
B Colpi long and narrow
- Lalongate pori
▪ Spheroidal, subprolate ~19  23 mm, relatively few, loosely distributed spines short and resembling equilateral triangles Sphaeranthus bullatus
▪ Spheroidal, circular ~28 mm, spines with broad, merged bases and acute tips, columellae extending into to broad part of spines Schkuhria pinnata
- Circular pori, grain spheroidal, circular ~25 mm, colpi long and wide Helichrysum citrispinum
- Lolongate pori, spheroidal, circular e triangular, ~28 mm, many relatively short spines broad based with acute tips Cineraria deltoidea
B Colpi long and wide (open)
- Lalongate pori, colpi without costa
▪ Colpi recessed, grain trilobate ~18 mm, exine inbetween spines distinctly granular Helichrysum forskhalii var. f.
▪ Colpi recessed, exine inbetween spines psilate/scabrate, grain trilobate ~21 mm, subprolate ~21  26 mm Helichrysum argyranthum
 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386 311

Table 3 (continued )

- Lalongate pori, colpi with costa

▪ Grain spheroidal, ~25 mm, pines with long (sometimes bent) acute tips/broad base, tip seems slightly set off the base Conyza vernonioides
▪ Grain spheroidal, slightly trilobate ~20 mm, spines with acute tips/broad base, tectum between spines is perforated to sub-psilate Conyza newii
▪ Grain spheroidal ~25 mm, spines with acute tips and broad base, more and smaller spines than in C. newii and C. vernonioides Conyza ruwenzoriensis
- Lolongate pori
▪ Colpi recessed, grain trilobate ~32 mm Senecio cyaneus
▪ Grains circular, spheroidal ~20 mm, tip often bent Dicrocephala integrifolia
▪ Grains circular, spheroidal ~25 mm, spines acute Helichrysum schimperi
- Circular pori, grain shperiodal ~30 mm Siegesbeckia abyssinica
- Pori inconspicuous, grain subprolate ~26  31 mm Spilanthes mauritiana
▪ Grains ≥ 34 mm
B Apertures undiscernible, grain spheroidal ~45 mm Haplocarpha rueppellii
B Colpi long and wide
- Pori lalongate, rectangular, grain spheroidal ~45 mm Tagetes minuta
- Pori circular-lolongate, grain subprolate 37  40 mm Crassocephalum bojeri
B Colpi long and narrow, circular pori, spines long and acute, ~41 mm Bidens pilosa
B Long colpi ± open, pori ± circular, short blunt spines with broad base, ~23 mm Stoebe kilimandscharica
B Colpi 1/3 to 2/3
- Colpi open, pori circular-lalongate, grain spheroidal ~40 mm Pluchea nitens
- Colpi narrow, pori lolongate, grain spheroidal ~54 mm Carduus keniensis
➢ Psilate
B Syncolpate
▪ Lalongate pori with vestibulum and thin annulus, grain triangular ~21 mm Syzygium guineense ssp. a.
▪ Lalongate pori protruding, grain subtriangular ~15 mm Solanum seaforthianum
B Colpi 2/3 of grain length
▪ Colpi constricted at equator, grain triangular-trilobate ~27 mm Indigofera atriceps subsp. k.
▪ Colpi recessed and narrow, pore lalongate, grain trilobate ~27 mm Trichilia emetica
▪ Colpi not recessed, grain not lobate
B Grain subprolate ~48  56 mm, elliptic broad pori Boswellia neglecta
B Grain spheroidal-subprolate ~20  22 mm, pore indistinct ± circular Rapanea melanophloeos
B Colpi >2/3 of grain length
▪ Lobate grain with deeply recessed colpi
B Colpi distinctly constricted at equator
- Colpi end blunt, grain subprolate ~23  28 mm Cassia spectabilis
- Colpi end acute, pore elliptic broad, grain spheroidal ~20  22 mm Dasylepis integra
B Colpi not or only slightly constricted at equator
- Grain subprolate-rectangular ~22  30 mm, thick exine smooth Euclea divinorum
- Grain subprolate-rhombic ~16  21 mm, large elliptic broad pori Crassula alsinoides
- Small grain spheroidal ~12 mm, with indistinct pori Nuxia congesta
- Double columella, pori lalongate, grain subprolate ~36  45 mm Gerbera viridifolia
- Grain subprolate, 36e42  50 mm, margo, indistinct pori lolongate Jacaranda mimosifolia
▪ Circular or triangular grain
B Grain subprolate and circular
- Pore circular and protruding, colpi narrow, grain ~27  36 mm Aningeria adolfi-friedericii
agg.
- Pore elliptic broad, colpi constricted at equator, grain ~18  23 mm Hypericum revolutum ssp. k.
B Grain subprolate and triangular
- Pore circular, colpi with broad margo, grain ~31  37 mm Cyphia glandulifera
B Grain spheroidal and circular or (sub)triangular
- Very large rectangular pori, grain triangular ~20 mm Desmodium repandum
- Pore long lalongate with vestibulum, grain subtriangular ~ 20 mm Solanum nigrum
➢ Scabrate
B Syncolpate, distinctly 3-armed, triangular concave
▪ Narrow colpi slightly recessed, arm length 20 mm, ~43 mm in polar view Englerina woodfordioides
▪ Arms ‘rectangular’ short, narrow colpi, ~37 mm in polar view Tapinanthus brunneus
B Colporate
▪ Grain circular and spheroidal/subprolate
B Pore without annulus
- Grain >30 mm
▪ Colpi (4/5) wide with thin margo and acute ends, constricted at equator,pori unclear, grain spheroidal, circular to slightly lobate Lobelia deckenii
~36 mm
▪ Colpi (4/5) wide with thick margo and obtuse ends, constricted at equator, pori unclear, subprolate ~31  39 mm, spheroidal to Lobelia holstii
slightly trilobate
- Grain <30 mm
▪ Lolongate, with fissured margins, subprolate 20  25 mm Alchemilla volkensii
▪ Lalongate pori, colpi (4/5) narrow with thin costa, grain ~20  25 mm Macaranga kilimandscharica
B Pore annulate
- Elliptic tall pori wider than long and narrow colpi, grain spheroidal ~27 mm Rumex bequaertii
- Large circular (Ø ~5 mm) to elliptic broad pori, grain spheroidal ~35 mm Cyphostemma
kilimandscharicum
- Pori elliptic broad, long (5/6) and narrow colpi, grain spheroidal ~18 mm Galiniera saxifraga
▪ Grain trilobate or triangular
B Colpi very short (<1/3)
- Colpi with thin margo, pori circular with operculum, grain trianguar ~ 24 mm Protea kilimandscharica
- Colpal area granular, pori inconsiciouos, grain triangular ~ 53 mm Turraea holstii
B Colpi long (>2/3)
(continued on next page)
312 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386

Table 3 (continued )

- Pori elliptic/rectangular broad, narrow colpi constricted at equator, Grain spheroidal ~19 mm Osyris lanceolata
➢ Reticulate
B Syncol(por)ate
▪ Pori circular to elliptic broad with annulus, colpi narrow, trilobate, prolate, ~31  56 mm Cissus oliveri
▪ Colpi with margo, tall elliptic pori with costa, circular ~44 mm, (sub)prolate ~44  60 mm Clutia abyssinica var. a.
▪ Syncolpate, wide colpi, margins granular, finely reticulate, subprolate, trilobate ~40 mm Markhamia lutea
▪ Syncolpate, sexine thinner around wide colpi, finely reticulate, circular, spheroidal/trilobate ~33 mm Tecoma stans
B 3-Colpate
▪ Colpi  1/2
B Colpi wide with acute ends, margins not clearly distinguishable, reticulum coarse and homobrochate, circular, spheroidal ~70 mm Psychotria petiginosa
B Colpi very short and narrow, angulaperturate, reticulum heterobrochate with large lumina, grain triangular convex to spheroidal Impatiens pseudoviola
~36 mm, circular
▪ Colpi > 1/2
B Colpi straight and narrow
- Colpi with thin margo, fine reticulum, subprolate ~25  31 mm, trilobate to circular Leucas volkensii
- Rough hexagonal reticulum, prolate ~24  44 mm, circular to triangular Blepharis maderaspatensis
- Recessed colpi narrow with thin margo, reticulum heterobrochate, (sub)prolate ~34  47 mm, trilobate-circular Leonotis mollissima
- High reticulum very coarse, grain trilobate and spheroidal ~23 mm Olea capensis ssp. h.
B Colpi distinctly wider at equator
- Colpi recessed and granular, reticulum homobrochate, subprolate/spheroidal ~30  35 mm, circular/trilobate ~33 mm Oxalis latifolia
- Colpi with very thin granulate margins, reticulum fine, subprolate ~20  23 mm, trilobate Caylusea abyssinica
- Colpi with smooth margins and obtuse ends, spheroidal/suboblate and trilobate/circular ~12 mm Trichocladus ellipticus
B 3-Colporate, colpi short (<2/3)
▪ Grain ± triangular
B Grain <40 mm
- Grain triangular ~ 25e28 mm, oblate, circular pore with vestibulum Apodytes dimidiata
- Grain convex triangular ~ 32 mm, rhombic, pore lalongate with operculum Hagenia abyssinica
- Grain slightly convex triangular ~49 mm, colpi arcuate with arci considerably thinner towards poles, pori lolongate with thick Rauvolfia caffra
annulus
B Grain >40 mm
- Colpi and pori large and unclear in shape, wide open in polar view, heterobrochate, spheroidal ~53 mm Psychotria fractinervata
- Colpi wide with acute ends, pori circular, grain spheroidal ~46 mm Erythrina burtii
▪ Grain circular to trilobate
B Pore rectangular broad
- Narrow colpi, grain circular to subprolate/rectangular tall ~21  28 mm Pentas lanceolata ssp. l. var. l.
- Colpi wider at equator, with acute ends, grain circular, spheroidal ~25 mm Pentodon pentandrus var. p.
B Other pore shape
- Small pori circular, heterobrochate, grain circular/spheroidal ~34 mm Psychotria capensis ssp. r.
- Macroreticulum heterobrochate, grain spheroidal/circular/trilobate ~56 mm Delonix regia
- Pori lolongate with thick annulus, colpi arcuate with arci considerably thinner towards poles grain ± circular ~49 mm, Rauvolfia caffra
B 3-Colporate, colpi 2/3
▪ Grains >50 mm and circular, spheroidal or subprolate
B Pore elliptic tall or lolongate
- Pori lolongate and sunken, colpi with irregular margins, subprolate ~69  78 mm,trilobate ~72 mm, grains mostly torn open along Convolvulus kilimandschari
colpi
- Colpi very wide, tapering at poles, colpoid streaks (intectate) very dictinct, pori elliptic tall, circular, spheroidal ~54 mm Caesalpinia decapetala
- Long (3/4) and narrow colpi, colpi tapering towards poles and with thicker margo at the equator, pori elliptic tall, subprolate Momordica boivinii
~61  72 mm, circular
B Pore elliptic broad or lalongate
- Colpi narrow with margo, pori inconsp., subprolate ~64  86 mm, circular Momordica foetida
- Colpi with large, elliptic broad pori, spheroidal, circular ~100 mm Luffa cylindrica
▪ Grains  50 mm
B Pore rhombic broad
- Pore rhombic with annulus, grain circular/spheroidal/subprolate ~25  29 mm Pavetta abyssinica var. a.
B Pori lalongate, elliptic, or rectangular broad
- Grain triangular, ~25 mm, oblate, rectangular pore protruding Scutia myrtina
- Grain circular/prolate
▪ Pori rectangular ~3  14 mm, colpi wide, prolate ~36  52 mm Grewia bicolor
▪ Lalongate pori with costa, narrow colpi, prolate ~15  31 mm Phyllanthus boehmii var.
boehmii
▪ Rectangular broad pori, colpi narrow, prolate/rectangular ~24  43 mm Triumfetta flavescens
▪ Rectangular broad/lalongate pori, narrow colpi constricted at equator, prolate e rhombic tall ~30  50 mm Triumfetta brachyceras
▪ Colpi narrow, slightly constricted at equator, large pori rectangular/elliptic, prolate to slightly rectangular ~17  29 mm Crotalaria axilliaris
- Grain circular and spheroidal/subprolate
▪ Lalongate pori, colpi relatively narrow, exine thickened at the pori, spheroidal to subprolate ~33  36 mm Euphorbia engleri
▪ Pori lalongate, colpus margins narrow, exine thickend at the pori, subprolate to spheroidal ~40  45 mm, circular to trilobate Euphorbia schimperiana var.
s.
▪ Colpi narrow without distinct margin, rectangular broad to lalongatepori, subprolate ~40  51 mm, circular (often folded) Grewia microcarpa
~50 mm
▪ Colpi slightly open constricted at equator, circular to oval pori, subprolate ~21  25 mm, circular to slightly trilobate ~21 mm Dalbergia lactea
▪ Colpi narrow, pori rectangular/elliptic, grain subprolate ~16  19 mm, slightly trilobate to circular ~19 mm Crotalaria keniensis
▪ Colpi narrow, pori rectangular to elliptic broad, grain subprolate to rectangular ~19  29 mm, spheroidal to slightly trilobate Crotalaria goodiformis
- Grain trilobate/(sub)prolate
▪ colpi wide at equator, pori large, (sub)prolate ~22  30 mm, Maerua grantii
▪ pori rectangular, colpi narrow, circular/trilobate, prolate ~18  26 mm Psychotria cyathicalyx
▪ pori lalongate, colpi narrow and straight, subprolate ~20  24 mm Toddalia asiatica
▪ colpi long (3/4) and narrow, pori rectangular, subprolate ~20  27 mm Cussonia arborea
 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386 313

Table 3 (continued )

▪ Pore butterfly shaped, grains trilobate/(sub)prolate, colpi narrow

- With thin margo, heterobrochate, prolate ~19  29 mm Schefflera volkensii
- Exine thick at poles, heterobrochate, prolate ~21  29 mm Schefflera myriantha
- Homobrochate, subprolate ~17  24 mm, slightly trilobate Polyscias fulva
B Pori lolongate, elliptic or rectangular tall
- Grain (sub)triangular
▪ Pori inconsp. lolongate, grain suboblate oblate, triangular ~42 mm, Strychnos scheffleri
▪ Colpi wide at the equator with acute ends, pori lolongate/elliptic tall, grain prolate ~19  29 mm, subtriangular Aeschynomene schimperi
▪ Very wide colpi with granular membraned margo, small lolongate poriwith vestibulum, grain triangular~27 mm, spheroidal- Pterolobium stellatum
subprolate
- Grain spheroidal to trilobate
▪ Colpi wide, pori elliptic tall, (sub)prolate ~20  28 mm, trilobate Dovyalis abyssinica
B Pori ± circular or inconspicuous
- Grain triangular
▪ Pore with lolongate annulus, thick margo, prolate ~35e40x50e65 mm Asystasia gangetica
▪ Colpi wide with margo, spheroidal ~28 mm Indigofera swaziensis subsp. s.
▪ Circular-squared pori with annulus, colpi narrow with margo, suboblate, triangular convex ~26 mm Hippocratea africana
- Grain circular to trilobate
▪ Large pori (Ø 8e9 mm) with annulus, colpi broader at equator, spheroidal ~36 mm Balanites maughamii ssp.
acuta
▪ Narrow colpi, pore small grain trilobate/prolate ~10x14 mmCleome hirta
▪ Colpi recessed with acute ends, slightly constricted at equator large circular pori, subprolate ~32  39 mm Eriosema montanum var. m.
▪ Colpi wide, pori inconspicuous, subprolate ~24  31 mm Calpurnia aurea
▪ Colpi wide with smooth margins, pori inconsipiouos, spheroidal ~19 mm Rawsonia lucida
▪ Colpi with smooth margins and obtuse ends, pori inconsipiouos, grain spheroidal/suboblate and trilobate/circular ~12 mm Trichocladus ellipticus
▪ Broad colpi with thin margo tapering towards poles, pori inconsipiouos,grain spheroidal and circular to trilobate ~21 mm Rourea thomsonii
➢ Striate/striato-reticulate
B Pori elliptic tall or circular
▪ Apertures weakly defined, granulate colpi 2/3, grain ± spheroidal ~26  30 mm, circular Lannea triphylla
▪ Large pori with annulus, colpi 4/5 narrow, spheroidal-trilobate, subprolate ~31  39 mm Balanites aegyptiaca
B Pori elliptic or rectangular broad or lalongate
▪ Grain trilobate and (sub)prolate, exine striate
B Narrow colpi, broad rectangular (~7  1.5 mm) pori/colpi transversales with costae, subprolate ~18  25 mm Rhus vulgaris
B Narrow colpi, pori elliptic broad, prolate ~9  13 mm Begonia johnstonii
▪ Grain circular/subtriangular and rhombic tall, exine reticu-striate
B Lalongate/rectangular pori narrow, colpi with margo, subprolat/rhombic tall ~21  29 mm, subtriangular Clausena anisata
B Pori rectangular broad ~3  10 mm, mesoporia slightly vaulted, rhombic tall ~30  35 mm, circular ~38 mm Ozoroa insignis
➢ Retipilat
B 3-colporate, square pori, colpi (2/3) narrow and tapering towards poles, trilobate, circular ~30 mm, sexine considerably thicker than Maytenus acuminata
nexine, retipilat/reticulate
➢ (Sub)Echinolophate
B Subechinolophate
▪ Grain circular and spheroidal ~47 mm, spines short with broad base, pore visibly lolongate/elliptic tall Vernonia syringifolia
▪ Grain slightly trilobate, recessed colpi, spines long and slender
B Spines often bent, ridges and lacunae clearly visible, of variable shape and size Erlangea calycina
B Separation of ridges and lacunae not always clear, spines longer than in Vernonia brachycalyx
B Echinlophate
▪ Short or long spines with broad (merged) base
B Short colpi and/or elliptic tall pori, lacunae regularly spaced, spines long, grain circular/spheroidal ~32 mm Tolpis capensis
B Colpi divided into three lacunae, aporal lacunae angular; paraporal lacunae pentagonal, spines short, grain circular/spheroidal Lactuca capensis
~40 mm
▪ Spines with slim base
B Grain circular, spheroidal >55 mm
- Ridges thick and high, lacunae angular but irregularly shaped, grain ~63 mm Bothriocline amplifolia
B Grain circular, spheroidal <55 mm
- Lacunae circular to hexagonal, ridges relatively low and thin, grain ~43 mm Bothriocline longipes
- Ridges very high, lacunae roundish and regularly spaced, spines acute, grain ~49 mm Vernonia lasiopus
- Lacunae roundish and regularly spaced, spines blunt, grain ~49 mm Vernonia colorata
- Colpi divided into three lacunae connected by narrow interlacunar gaps, abporal lacunae large, angular and broader towards the Crepis oliveriana
poles, grain ~42 mm
➢ Psilolophate
B 3-colporate, ridges high, circular pori, circular grain ~16 mm (incl. sculpture) Hirpicium diffusum
➢ Verrucate
B 3-colpate, colpi long and wide, (sub)prolate ~50  70 mm, circular Plumbago dawei
B 3-colporate, syncolpate, colpi recessed, pori elliptic tall, triangular ~29 mm, suboblate Garcinia volkensii
15) 4-aperturate
➢ Colpate, reticulate
B Grain circular, spheroidal
▪ Colpi short ½, narrow, grain ~44 mm Chassalia kenyensis
▪ Colpi very short 1/5, narrow, grain ~47 mm Chassalia parviflora
B Grain tetralobate or rectangular broad
▪ Colpi 3/5, grain tetralobate ~38 mm and spheroidal/subprolate, sexine > nexine Jasminum abyssinicum
▪ Colpi very short (max 1/6), grain rectangular broad ~29  47 mm, oblate Impatiens walleriana
➢ Colporate
B Echinate, very short oval colpi, lalongate/elliptic broad pori, circular/spheroidal ~36 mm Tridax procumbens
B Psilate, colpi long (>3/4) and narrow
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Table 3 (continued )

▪ Pori elliptic broad with annulus, grain rectangular tall ~8  15 mm, quadrangular Lithospermum afromontanum
▪ Circular pori protruding, grain subprolate-rectangular, circular, ~27  36 mm Aningeria adolfi-friedericii
agg.
B Scabrate, colpi short (<1/2)
▪ Pori broad rectangular, mesopori vaulted and granular, spheroidal, circular ~40 mm Tabernaemontana stapfiana
▪ Elliptic broad/lalongate pori with annulus, circular/quadrangular ~21 mm, spheroidal Lepidotrichilia volkensii
▪ One ulceroid aperture at the thick end and 3 indistinct lateral elliptic poroid regions, thin exine, concial, longest axis ~53 mm Carex conferta
➢ Porate
B Granulate/bacculate, circular/elliptic pore with thick annulus, circular, spheroidal, ~25 mm Celtis gomphophylla
B Scabrate, exine thicker around pori, vestibulum, quadrangular ~34 mm, oblate Allophylus ferrugineus
B Finely reticulate, 3 (4) porate, large elliptic pori, grain triangular ~ 44 mm, (sub)oblate Paullinia pinnata
16) 5-aperturate
➢ Heterocolporate
B 3-pseudocolpato-2-colporate, psilate, pori circular annulate, spheroidal, circular ~34 mm Saba comorensis
B One ulceroid aperture at the thick end, 4 lateral elongate poroid regions, coarsely scabrate to verrucate, spheroidal to ovoid ~35 mm, Cyperus amauropus
circular, heteropolar
B One ulceroid aperture at the thick end and 4 indistinct lateral elongate poroid regions, ovoid ~65 mm, heteropolar Carex vallis-rosetto
➢ Colporate
B Scabrate, short colpi, elliptic broad/lalongate pori with annulus, ± circular ~21 mm, spheroidal Lepidotrichilia volkensii
➢ Porate
B Reticulate, pori Ø ~5 mm ± circular with annulus (endocosta), circular, spheroidal ~22 mm Mussaenda microdonta
B Echinate, por elongate but often obscure, circular, spheroidal ~104 mm Sida ovata
17) 6-aperturate
➢ Colporate
B Elliptic pori, narrow colpi of 1/3 length, quincuadrangular ~33  33 mm Basella alba
➢ Heterocolporate (3-colpori-3-pseudocolporate)
B Psilate/scabrate/granulate
▪ Pori ± circular Ø ~4.5 mm, psilate/scabrate. subprolate ~23  30 mm, hexalobate Combretum apiculatum
▪ Colpi constricted at equator, pori elliptic/rectangular tall, scabrate/granulate, rectangular tall to prolate ~25  34 mm, hexalobate Combretum psidioides subsp.
p.
▪ Pori elliptic tall with annulus, psilate, rectangular tall ~23  30 mm, circular/trilobate to slightly hexalobate Heliotropium rariflorum ssp.
h.
B Reticulate
▪ Colpi long, narrow, constricted at the equator, pseudocolpi shorter, faint and wider, pori elliptic broad, prolate ~20  30 mm, triangular to Ehretia bakeri
slightly hexalobate
➢ Heterocolporate (2-colpori-4-pseudocolpate)
B Colpi long and narrow, pore circular, grain reticulate, subprolate 42  32 mm Justicia asystasioides
➢ Colpate
B Circular, spheroidal, reticulate
▪ Very narrow colpi (3/5 to 4/5) with acute ends, coarsely reticulate, grain ~21 mm Kohautia coccinea
B Subprolate to prolate, spheroidal, reticulate, colpi long
▪ Exine distinctly thicker at poles, reticulate with very regular sized lumina, at poles reticupilat, prolate ~42  52 mm, circular ~42 mm Plectranthus sylvestris
▪ Exine thicker at poles, reticulate with distinctly smaller lumina in polar area, prolate ~33  45 mm, circular ~40 mm Plectranthus alboviolaceus
▪ Muri with open ends in the equator area, partly very long lumina, smaller in polar area and generally larger than in P. alboviolaceus, Plectranthus laxiflorus
prolate ~34  42 mm, circular ~41 mm
18) Polyaperturate (>6), peri- and stephanoaperturate
➢ Pericolpate
B Microechinate
▪ Colpi short and recessed with irregular margins, circular, spheroidal ~43 mm Ranunculus oreophytus
B Psilate, scabrate, verrucate
▪ One ulceroid aperture at the thick end, 5e6 distinct lateral mostly very long conical colpoid regions, conical, longest axis ~35 mm, Cyperus rigidifolius
heteropolar Richardia scabra
➢ Periporate, grain circular and spheroidal
B Echinate
▪ Columella rod-like, elongated beneath the bases of spines
 Many circular pori Ø ~4 mm, ± annulate, sparsely distributed spines ~10 mm, straight, slender, with blunt apices, grain ~90 mm Hibiscus meyeri
 Many circular pori Ø < 2 mm, tectum micro verrucate/granulate, spines clubbed ~6e10 mm with bulgy base ± acute apices, grain Hibiscus adoensis
~100 mm
 Circular pori Ø 4e5 mm, wide annulus, parsely distributed spines ~ 15e20 mm with subacute tips, grain ~110 mm Pavonia elegans
▪ Columella not visible/not rod-like
 Many circular pori Ø ~ 2 mm, spines ~5 mm, grain ~71 mm Astripomoea hyoscyamoides
subsp.h.
B Psilate, scabrate
▪ Tectum perforate with a few microspines
 Circular pori Ø 4e5 mm with irregular border, mesorporia flat, grain ~24 mm Celosia schweinfurthiana
 Circular pori Ø 2e4 mm with thin annulus, irregular border, grain ~22 mm Achyranthes aspera
 Circular sunken pori Ø 1e2 mm with sharp border, grain ~25 mm Amaranthus hybridus
▪ Tectum complete without microspines, scabrate
 Circular pori Ø 1.5 mm, exine thick, finely microreticulate, grain ~21 mm Gnidia apiculata
 Few sunken, roundish pori Ø 3.5e5 mm with irregular margins, grain ~19 mm Thalictrum rhynchocarpum
B Reticulate
▪ Microreticulate, heterobrochate, lumen smaller around pori
 9e15 circular pori Ø ~5 mm with annulus, grain ~30 mm Cerastium indicum
 15e20 circular pori Ø ~3 mm with annulus, grain ~33 mm Uebelinia rotundifolia
▪ (Coarsely) reticulate, lumina larger than above, exine thick
 Inconspicuous circular pori, coarsely reticulate with high muri, grain ~53 mm Polygonum salicifolium
 Circular pori small Ø ~2 mm (size of lumina), grain ~34 mm Gnidia glauca
 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
Table 3 (continued )
 Circular pori small Ø ~2 mm, distinctly > (about double) lumina, grain ~31 mm
B Verrucate
▪ > 40 small circular pori Ø ~1 mm with narrow annulus, grain ~32 mm
▪ < 10 roundish pori, sunken with granulate margins, grain ~24 mm
B Bacculate
▪ Stephanocolp(or?)ate, 21 very short colpi, bacculate, circular, spheroidal ~86 mm
19) Spiraperturate
➢ One spiral apertures, exine psilate/scabrate, circular, spheroidal ~64 mm
5. Pollen and spore atlas
The plant species covered in this atlas are subdivided into spores
(Pteridophytes) (Table 1), and pollen grains (Angiosperms and
Gymnosperms) (Table 2). The plant families are sorted alphabeti-
cally; the respective species are sorted also alphabetically by genus.
In case latest taxonomic research has shown a different affiliation of
species (as per Tropicos.org, 2015; Stevens, 2015), this is indicated
in brackets behind the species name. All families are introduced
with general information on the habitus, and distribution of the
plants, and continued with the spore/pollen morphology typical for
all genera included in the families following Erdtmann (1952, 1957;
1965) and Tryon and Lugardon (1991), and more recent literature
on pollen morphology on plant family level. The morphology of
pollen and spores of the individual species presented in this atlas is
then described in more detail. The identification key presented in
Table 3 is compiled based on distinctly discriminating morpho-
logical features of the different pollen types.
5.1. Spore types
5.1.1. Actiniopteridaceae
Terrestrial ferns; rhizome shortly creeping, densely covered
with scales as are the bases of stipes. The family includes a single
genus extending from Africa to Arabia, Mascarene Islands, Comoro
Islands, Madagascar, India and Sri Lanka (Verdcourt, 1999a).
5.1.1.1. Spore morphology. Trilete spores.
Actiniopteris dimorpha Pic. Serm (Plate 1, 1e2). (Pteridaceae).
Shape: triangular (convex) ~55e62 mm.
Apertures: trilete, laesura 2/3 to 3/4 the radius.
Surface: distal face densly beset with irregular verrucae;
rugulate.
Structure: exine is thicker at distal face and thinner in proximal
face; probably with perine.
Habitus: fronds dimorphic, the fertile about twice as long as the
sterile and less numerous. Fertile fronds (8-) 15e18 (27) cm long.
Habitat: inselbergs and rock domes, in shaded crevices, usually
dry forest of Commiphora, Lannea etc with Xerophyta, Jatropha,
Anthericum etc.; also Julbernandia globiflora woodland;
550e1400 m.
At Kilimanjaro sun-exposed cliffs in the savanna and sub-
montane cultivation zone, growing together with Actiniopteris
raddiata, A. semiflabellata, Cyperus amauropus, Crassula volkensii and
Xerophyta spekei, from 800 to 1500 m.
Distribution: U 3; K 1, 2, 4, 7; T 2e8; DR CONGO, Burundi, Sudan,
Ethiopia, Somalia, Zambia, Malawi, Mozamique, Zimbabwe,
Madagascar, Comoro Is and Re union. At Kilimanjaro very rare.
5.1.2. Adiantaceae
Terrestrial or rupestral ferns of small to large (over 2 m high)
size. The family is pantropical with extensions northward to
Newfoundland, Alaska, and northeastern Asia, and southward
to southern South America, southern Africa, and New Zealand
315
Gnidia subcordata
Chenopodium ambrosioides
Plantago palmata
Richardia scabra
Thunbergia alata
(Tryon et al., 1990).
5.1.2.1. Spore morphology. Spores trilete, tetrahedral-globose with
prominent angles, more or less depressed between the aperture
arms, irregularly tuberculate or rugose below irregular sheaths,
without an equatorial flange (Tryon et al., 1990).
Adiantum capillus-veneris L (Plate 1, 3e5). (Pteridaceae).
Shape: triangular ~50e60 mm, the angles usually prolonged.
Aperture: trilete, laesura 3/4 the radius.
Suface: irregular sheaths over a rugulate surface.
Structure: exospore usually plain; perispore of two layers, the
outer forms the rugulate surface or three complex layers form the
largest part of the sporoderm.
Habitus: rhizome creeping, short, ± coarse; fronds 5e50, clus-
tered, erect or spreading or hanging, 5e50 (90?) cm tall.
Habitat: roadbanks, riverbanks, wet rocks, caves etc., mostly in
forest or at least in shade; 200e3000 m. At Kilimanjaro on wet
shaded rocks in riverine forests from the savanna to the lower
montane zone (800e1950 m), growing together with Selaginella
abyssinica and Pilea rivularis.
Distribution: U 1e4; K 1, 3, 4, 6, 7; T 1e3, 4, 5e8; cosmopolitan.
At Kilimanjaro scattered.
Adiantum incisum Forssk (Plate 1, 6e7). (Pteridaceae).
Shape: straight to concave triangular ~31 mm, the angles usually
prolonged.
Aperture: trilete, laesura 3/4 the radius with distinctly darker
costa.
Suface: loosely and finely rugulate.
Structure: exospore usually plain; perispore of two layers, the
outer forms the rugulate surface or three complex layers form the
largest part of the sporoderm.
Habitus: rhizome short, erect. Fronds tufted, erect or decum-
bent, 10e40 cm long.
Habitat: pathsides, river banks, under and between rocks, bases
of trees, walls etc. in shady places, often riverine forest; 0e1650
(1800?) m. At Kilimanjaro at the colline savanna foothills in semi-
shaded rock crevices inside riverine forests, in the submontane
cultivation zone on humid embankments of irrigation funnels and
road ditches in the same habitat as Adiantum poiretii, from 800 to
1600 m.
Distribution: U 1e4; K 1, 2, 4, 6, 7; T 1e3, 5e8; Z; widespread in
tropical Africa; South Africa; Yemen. At Kilimanjaro scattered.
Adiantum poiretii Wikstr. var poiretii (Plate 1, 8e10)
(Pteridaceae).
Shape: triangular ~49 mm, the angles usually prolonged.
Aperture: trilete, laesura 3/4 the radius with smooth costa.
Suface: granulate to rugulate.
Structure: exospore usually plain; perispore of two layers, the
outer forms the rugulate surface or three complex layers form the
largest part of the sporoderm.
Habitus: rhizome creeping, slender but with thicker branches.
Fronds closely spaced, erect, (10-) 20e60 (100) cm tall.
Habitat: streamsides, rocky gorges, waterfalls, swampy areas,
sometimes on bare earth, rock-clefts, mostly in forested areas,
316 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386

Plate 1. ACTINIOPTERIDACEAE: 1e2 Actiniopteris dimorpha; ADIANTACEAE (Pteridaceae): 3e5 Adiantum capillus-veneris, 6e7 Adiantum incisum, 8e10 Adiantum poiretii var. poiretii,
11e12 Coniogramme africana, 13e15 Pellaea dura var. dura, 16e18 Pityrogramma aurantiaca.

montane forest including forest/bamboo boundary; 1150e2700 m. between aperture arms.

At Kilimanjaro shaded streamsides and crevices in riverine forests
in similar habitats as A. capillus-veneris, humid shaded embank-
ments of irrigation funnels and road ditches in the Chagga home-
gardens and here growing together with A. hispidulum, A.
raddianum, A. incisum and Pteris dentata, from 800 to 2300 m.
Distribution: U 1e4; K 1e6; T 2e4, 6e8; throughout tropical and
temperate Africa; Madagascar; Mascarene Is.; India; Tristan de
Cunha; Central and S America from Mexico to Uruguay. At Kili-
manjaro widespread.
Coniogramme africana Hieron (Plate 1, 11e12). (Pteridaceae).
Shape: triangular ~46 mm with prominent angles, depressed
Aperture: trilete, the laesura 3/4 the radius, leasura very narrow.
Surface: irregularly papillate.
Habitus: rhizome creeping, up to 8 mm in diameter. Fronds
spaced, 1e4 cm apart, herbaceous, 0.5e2 m tall.
Habitat: lower montane forest and rain-forest with Entandro-
phragma, Parinari, Bridelia, Macaranga, Allanblackia, Chrysophyllum
etc. by rivers and waterfalls; 1000e2250 m. At Kilimanjaro in lower
montane ravine forests with Entandrophragma, Newtonia, Strom-
bosia and Garcinia, from 1300 to 1900 m.
Distribution: U 2, 4; K 3e5; T 2, 3, 6, 7; widespread in tropical
Africa from Ghana, Liberia, Nigeria, Cameroon, Bioko, Sao Tome  to
 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
DR Congo-Kinshasa, Ethiopia and Malawi. At Kilimanjaro very rare
and restricted to the wet southern slope.
Pellaea dura (Willd.) Hook var dura (Plate 1, 13e15)
(Pteridaceae).
Shape: triangular ~ 54 mm with prominent angles, depressed
between aperture arms.
Aperture: trilete, the laesura 3/4 the radius, leasura opened with
costa.
Surface: psilate.
Habitus: rhizome shortly creeping or ascending. Fronds 3e15,
tufted, 7e50 cm tall.
Habitat: grassland, Brachystegia woodland, Xeroderris-Afzelia-
Pterocarpus woodland, crevices in rocky slopes; 600e1800 m. At
Kilimanjaro particulary in anthropogenic dry grassland of the
submontane zone with Hyparrhenia rufa, Satureia abyssinica, Aris-
tida adoensis and Crepis carbonaria, from 900 to 1600 m.
Distribution: U 2; T 2, 3, 6, 7; Angola, Zambia, Malawi,
Mozambique, Zimbabwe, Nigeria, Central African Republic,
Cameroon, Gabon, DR Congo-Kinshasa, Burundi, Sudan, Ethiopia,
South Africa, Madagascar, Re union. At Kilimanjaro rare.
Pityrogramma aurantiaca (Hieron.) C.Chr (Plate 1, 16e18).
(Pteridaceae).
Shape: triangular ~54 mm, the distal face sometimes
hemispherical.
Aperture: trilete, the laesura 2/3 to 3/4 the radius, often with
ridges parallel to aperture.
Surface: usually coarsely tuberculate or rugate with prominent
equatorial flange, sometimes granulate without rugae or flange.
Structure: exospore of two layers, the contours formed by the
outer one; perispore two layered, often with papillate surface.
Habitus: rhizome short to long-creeping. Fronds closely spaced,
erect or ± scrambling.
Habitat: moorland with Erica etc., upper part of montane forest,
among rocks in volcanic craters; (1460-) 2000e3500 m. At Kili-
manjaro mainly in upper montane forests with Podocarpus, Hage-
nia, Erica excelsa, also in subalpine Festuca obturbans tussock
grasslands, from 2400 to 3200.
Distribution: U 2; K 4; T 2, 3, 6, 7; DR Congo-Kinshasa, Rwanda,
Burundi, Malawi. At Kilimanjaro scattered.
5.1.3. Aspleniaceae
Terrestrial, epilithic, or epiphytic ferns of small to medium rarely
to large size. The more than 700 species of Asplenium are distrib-
uted all over the world. The majority are in the (sub)humid tropics
and south-temperate regions (Kramer and Viane, 1990).
5.1.3.1. Spore morphology. Spores monolete, usually ellipsoid; per-
ispore prominent, consisting of a thin basal layer (inner perispore)
pressed against the exospores, this inner perispore bearing rod-like
(sometimes forking) structure (middle perispore) sustaining the
outer perisporal layer. Middle and outer variously developed, the
outer layer often (partly) perforated and/or thrown into variously
shaped ridges or into spines (Kramer and Viane, 1990).
Asplenium adiantum-nigrum L (Plate 2, 1e2).
Shape: ellipsoidal ~50 mm, rarely somewhat spheroidal.
Aperture: monolete, 2/3 to 3/4 the length of the spore.
Surface: folds usually prominent forming long wings (alate) or
cristate, echinate, or reticulate; the areoles between folds plain or
usually echinate, fenestrate, or reticulate.
Structure: exospore plain; perispore cavate, usually with pillars,
and long laminae in the inner part.
Habitus: rhizome creeping, to 4 mm thick. Fronds tufted.
Habitat: moist lava or rock, roadside, heath scrub, moorland;
1650e4300 m. At Kilimanjaro bimodal distribution below and
above the forest belt: from 1300 to 1800 and from 2900 to 4400 m.
317
Below the forest belt growing together with Pellaea viridis and
Adiantum raddianum on road and path embankments and hedges
inside Chagga homegardens as well as in Hyparrhenia rufa grass-
lands. Above the forest belt in alpine rock vegetation together with
Asplenium uhligii, A. praegracile, Melpomene flabelliformis and Arabis
alpina.
Distribution: K 1, 3, 4, 6, 7; T2; Cameroon; Chad; South Africa;
Morocco; Algeria; Europe; N America; Hawaii. At Kilimanjaro
scattered.
5.1.4. Cyatheaceae
Large to very large, less often medium-sized, terrestrial ferns
with short to usually long, erect, slender to robust, apically sclay
stem; the species are concentrated in the tropics where they are
most numerous in montane to (locally) alpine vegetation (Kramer,
1990a). Continental Africa has 14 species (Holttum, 1981).
5.1.4.1. Spore morphology. Spores trilete, heteropolar, radially
symmetric, scabrate, granulate, fossulate, perforate; laesura nar-
row, reaching the equator or extending 2/3 of the radius; laesura
bordered by broad margo (Roubik and Moreno, 1991).
Cyathea humilis Hier. in Engl. var humilis (Plate 2, 3e5).
Shape: triangular to triangular concave ~48e50 mm often with
prominent angles.
Aperture: trilete; the laesura 3/4 the radius, broad margo of
typically rounded shape.
Surface: mostly strongly granular.
Structure: Exospore forms the major contours, usually pitted,
sometimes lobed; thin perispore, often of two layers with rodlets
lining and spanning the pits.
Habitus: caudex absent or up to 2.5 m high and to 4 cm in
diameter.
Habitat: moist forest, often near streams, (?500-) 900e2250 m.
At Kilimanjaro restricted to streamsides in moist montane Ocotea
forest, from 2000 to 2200 m.
Distribution: K 4. 7: T 2, 3, 6, 7; Sudan. At Kilimanjaro extremely
rare and only on the wet southern slope.
Cyathea manniana Hook (Plate 2, 6e7).
Shape: strongly concave triangular ~40e42 mm often with
prominent angles.
Aperture: trilete; the laesura 3/4 the radius, broad margo of
typically rounded shape.
Surface: psilate to scabrate.
Structure: exospore forms the major contours, usually pitted,
sometimes lobed; thin perispore, often of two layers with rodlets
lining and spanning the pits.
Habitus: trunk 0.3e9 m tall, 10e15 cm in diameter. Fronds 5e10
in number, to 4 m long.
Habitat: gregarious, in forest along streams or adjacent to
springs, often forming a secondary canopy in high rainfall areas,
occasionally going up into the tree heath zone, 850e2700 m. At
Kilimanjaro mainly in moist montane Ocotea forest, in the sub-
montane zone only in riverineforest, from 1200 to 2900 m.
Distribution: U 2e4; K 1e7; T 2e8; widespread in tropical Africa
from Liberia to Kenya and south to Mozambique and Zimbabwe. At
Kilimanjaro widespread and often dominant, but nearly exclusively
on the wet southern slope.
5.1.5. Dryopteridaceae
Terrestrial or epilithic, very rarely epiphytic ferns of very
different size. The family is of cosmopolitan distribution (Kramer
et al., 1990).
5.1.5.1. Spore morphology. Spores without exception monolete,
with prominent perispore.
318 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386

Plate 2. ASPLENIACEAE: 1e2 Asplenium adiantum-nigrum, CYATHEACEAE: 3e5 Cyathea humilis, 6e7 Cyathea manniana, DRYOPTERIDACEAE: 8e10 Arachniodes webbiana subsp.
foliosa, 11e12 Didymochlaena truncatula, GLEICHENIACEAE: 13 Dicranopteris linearis, GRAMMITIDACEAE: 14e16 Xiphopteris flabelliformis, HYMENOPHYLLACEAE: 17e18 Hymeno-
phyllum capillare, 19e20 Trichomanes melanotrichum, LOMARIOPSIDACEAE: 21e23 Elaphoglossum acrostichoides, LYCOPODIACEAE: 24e25 Huperzia dacrydioides, 26e27 Huperzia
ophioglossoides, 28e29 Huperzia saururus, 30e32 Lycopodium clavatum, Marattiaceae: 33e34 Marattia fraxinea, OPHIOGLOSSACEAE: 35e37 Ophioglossum vulgatum subsp. kili-
mandscharicum, POLYPODIACEAE: 38e39 Loxogramme lanceolata.

Arachniodes webbiana subsp. foliosa (C. Chr.) Gibby, Rasbach,
Reichst., Widen & Viane (Plate 2, 8e10) SYN Dryopteris foliosa.
Shape: ellipsoidal ~25  38 mm.
Aperture: monolete, 2/3 to 3/4 the length of the spore.
Surface: echinulate.
Structure: exospore plain; perispore cavate, the inner part
irregularly papillate, sometimes pillared.
Habitus: rhizome creeping, 60e90 cm long, up to 7 mm in
diameter. Fronds spaced, arching, firmly herbaceous, 35e90 cm tall.
Habitat: montane and intermediate forest with Podocarpus,
bamboo, etc., stream gulleys, high rainfall areas; (?1250-)
1380e2600 (3130 Mt Elgon) m. At Kilimanjaro mainly in the
lower and middle montane zone in moist Ocotea, Cassipourea and
riverine forests, from 1300 to 3000 m.
Distribution: U 2, 3; K 1, 3e7; T 2, 3, 6; Rwanda, Malawi,
Zimbabwe, South Africa. At Kilimanjaro widespread but in low in-
dividual numbers.
Didymochlaena truncatula (Sw.) J.Sm (Plate 2, 11e12).
Shape ellipsoidal ~64  73 mm.
Aperture: monolete, 1/4 to 1/2 the length of the spore.
Surface: coarse, inflated, with compressed, often perforate,
winglike folds.
Structure: cavate perispore, the inner part papillate.
Habitus: terrestrial; rhizome up to 2.5 cm in diameter, erect,
forming a short woody caudex up to 20 cm high and 15 cm in
diameter. Fronds tufted, 0.7e2.1 m tall but up to 3.6 m long, arching,
firmly herbaceous.
Habitat: evergreen forest including mist and rain forest;
1000e2250 m. At Kilimanjaro mainly in submontane-lower
montane riverine and gorge forests, together with Marattia frax-
inea and Dryathyrium boryanum, from 1300 to 2200 m.
Distribution: U 2e4; K 4e7; T 2e4, 6e8; Bioko, Sa ~o Tome ,
Cameroon, DR Congo, Burundi, Rwanda, Ethopia, Angola, Malawi,
Mozambique, Zimbabwe, South Africa, pantropical. At Kilimanjaro
 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
scattered and only on the wet southern slope.
5.1.6. Gleicheniaceae
Terrestrial ferns of rather small to very large size with slender, at
least partly creeping, protostelic (with one exception) stem bearing
scales and/or hairs (Kramer, 1990b).
5.1.6.1. Spore morphology. Spores monolete; heteropolar-bilateral;
sclerine psilate; surface rugulose, low ridges forming a nearly
reticulate surface; laesurae curved, 2/3 length of spore; narrow;
acute apex; laesura with broad margo; amb elliptical, concave-
convex; perispore thin (Roubik and Moreno, 1991).
Dicranopteris linearis (Burm. f.) Underw (Plate 2, 13).
Shape: (mostly straight) triangular ~50 mm, with prolonged
angles.
Aperture: trilete, the laesura 2/3 to 3/4 the radius of the spore,
very distinct and broad margo.
Surface: low, usually foveolate and granulate.
Structure: exospore of three layers in the apertural region;
perispore thin, rarely with short rodlets.
Habitus: rhizome 1e5 mm in diameter. Fronds (0.1-) 1e1.8 m
long, spaced 6e20 cm apart with 1e3 levels of false dichotomy in
each lateral branch-system arising from each side of the terminal
bud.
Habitat: forest including rain-forest and bamboo-Juniperus,
forest fringes, rocky banks by streams, also by stream jets and fu-
maroles with Tarchonanthus, Dodonaea, Dissotis, etc, 900e2600 m
and also coastal down to 9 m. At Kilimanjaro a pioneer species on
disturbed ground on eroded soils, landslides, recent road em-
bankments and forest edges growing together with Lycopodiella
cernua from 1200 to 1800 m.
Distribution: U 2, 4; K 1, 3, 4, 7; T 2, 3, 6, 7; Z; P; Sierra Leone,
Liberia, Ivory Coast, Ghana, Nigeria, Cameroon, Bioko, Principe, Sa ~o
Tome  Gabon, Central African Republic, DR Congo, Rwanda, Ethiopia,
Angola, Zambia, Malawi, Mozambique, Zimbabwe, South Africa,
Madagascar, Comoro Is., Seychelles, Mauritius, Re union, Asia,
Australia, Polynesia. At Kilimanjaro rare, but locally common and in
species-rich colonies.
5.1.7. Grammitidaceae
Epiphytic, sometimes terrestrial or rupestral ferns, with creep-
ing to erect, simple or ± branched solenostelic stem usually bearing
scales, occasionally glabrous. Most species occur in tropical
montane primary rainforest, but some grow in tropical lowland
primary rainforests and others are found in lowland to montane
temperate rainforest or subalpine scrubland and alpine commu-
nities (Parris, 1990).
5.1.7.1. Spore morphology. Spores thin walled, usually globose to
tetrahedral with trilete laesura, papillate with scattered spherical
deposit, sometimes somewhat verrucate, somewhat echinate in
Calymmodon (Parris, 1990).
Xiphopteris flabelliformis (Poir.) Schelpe (Plate 2, 14e16) SYN
Melpomene flablliformis (Polypodiaceae).
Shape: triangular convex to spheroidal ~42 mm.
Apertures: trilete, laesura nearly equal to radius, wider in the
center.
Surface: loosely verrucate with different sized verrucae.
Structure: probably without perine.
Habitus: rhizome long-creeping, sometimes branched.
Habitat: usually a low epiphyte, often in moss cushions on tree
trunks and branches, in giant heath or Podocarpus/Hagenia forest to
Hagenia woodland, afroalpine bush, bamboo zone, sometimes on
rocks of old lava flows; 1000e4200 m. At Kilimanjaro mainly
epiphytic in upper montane Podocarpus, Hagenia and Erica excelsa
319
forest and epilithic in respective riverine forests together with
Elaphoglossum subcinnamomeum, Cheilanthes farinosa and Cys-
topteris nivalis; above the forest belt in the (sub)alpine zone on
rocks in the same habitat as Asplenium adiantum-nigrum, from 1900
to 4020 m.
Distribution: U 2, 3; K 3, 4; T 2e4, 6, 7; Cameroon, Bioko, DR
Congo, Rwanda, Ethiopia, Malawi, Mozambique, Zimbabwe, South
Africa, Madagascar, Mauritius, Re union. At Kilimanjaro scattered,
mainly on the southern slope.
5.1.8. Hymenophyllaceae
Epiphytic, saxicolous, or terrestrial fern of small to medium size;
stem creeping, or less often erect, with a so-called Hymeno-
phyllacean protostele, bearing only hairs (Iwatsuki, 1990). A nearly
worldwide family of eight genera especially of humid regions
(Tryon et al., 1990).
5.1.8.1. Spore morphology. Spore trilete; radially-symmetric; scler-
ine psilate, granulate, verrucate, foveolate; laesura reaching equa-
tor, narrow, irregular; laesura often with margo; amb subangular-
biconvex to circular (Roubik and Moreno, 1991).
Hymenophyllum capillare Desv (Plate 2, 17e18).
Shape: triangular convex ~55 mm.
Apertures: trilete, laesura 2/3 to nearly equal to radius, wider in
the center.
Surface: low pappilate, rarely echinate.
Structure: exoxpore three layers, the outer forming the surface
contours, perispore very thin.
Habitus: pendulous epiphyte or hanging from rocks; rhizome
long-creeping, filiform. Fronds 1e5 cm apart, pendulous.
Habitat: on treetrunks, less often on mossy branches and moist
cliff faces, within moist forest, montane evergreen forest and tree
heath, may be mat-forming and often locally common, (1000-)
1650e3150 m. At Kilimanjaro in moist lower-upper montane
Ocotea, Podocarpus and riverine forests, mainly epiphytic on Ocotea
usambarensis together with Hymenophyllum kuhnii, H. splendidum
and H. tunbrigense, also epilithic in upper montane riverine forests
together with Elaphoglossum subcinnamomeum, Cheilanthes far-
inosa and Cystopteris nivalis in the same habitat as Melpomene fla-
belliformis, from 1700 to 3300 m.
Distribution: U 2, K 3e5, 7; T 2, 3, 6, 7; widespread in tropical
Africa from Ghana to South Africa, Mascareignes, Madagascar. At
Kilimanjaro scattered, nearly exlusively on the southern slope.
Trichomanes melanotrichum Schltdl. (Plate 2, 19e20).
Shape: spheroidal ~52e55 mm.
Aperture: trilete, laesura 3/4 to nearly equal the radius, often
wiggeling, thin margo.
Surface: usually shallowly papillate to somewhat bacculate, or
echinate.
Structure: exospore of three layers forming the contours; peri-
spore not observed.
Habitus: lithophyte or epiphyte. Rhizome creeping, branched,
filiform. Fronds sometimes pendulous, but more often erect.
Habitat: moist forest and then in moist and shady sites, occa-
sionally in riverine or drier type forest, on tree-trunks, moist rock-
faces, mossy rocks, rarely on moist earth banks, may be locally
common, (200-) 750e2650 m. At Kilimanjaro mainly epiphytic in
lower-middle montane Ocotea forests, in lower elevations on rocks
in Mitragyna and Newtonia riverine and gorge forests, from 1200 to
2800 m.
Distribution: U 1e4; K 1e7; T 2e4, 6e8; Sierra Leone to Ethiopia
and south to South Africa, Madagascar, Mascarene Is. At Kilimanjaro
scattered.
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5.1.9. Lomariopsidaceae
Terrestrial, epilithic, scandent, or epiphytic, small to rather large
ferns with short-to long-creeping (or -scandent), less often erect
stem bearing basifixed or narrowly peltate, clathrate or non-
clathrate scales that are often marginally toothed or ciliate
(Kramer, 1990c).
5.1.9.1. Spore morphology. Spores monolete, mostly with very
prominent, folded perispore. The spores are monolete, ellipsoidal,
usually with prominent, wing-like folds or with reticulate surface.
Elaphoglossum acrostichoides (Hook. & Grev.) Schelpe (Plate 2,
21e23) (Dryopteridaceae).
Shape: ellipsoidal ~30  42 mm.
Aperture: monolete, 1/2 to 2/3 the length.
Suface: inflated folds, winged, cristate.
Structure: exospore plain; perispore a single cavate layer, often
with abundant pillars.
Habitus: rhizome moderately creeping, ± 4 mm diameter;
sterile fronds 4e10 mm distant, (6-) 20e50 cm long.
Habitat: epiphytic, less commonly on rocks or soil, in moist
montane forests; 1900e2700 m. At Kilimanjaro mainly epiphytic in
lower-upper montane Ocotea, Podocarpus and Erica excelsa forest
from 1500 to 2800 m.
Distribution: U 2; K 3e5; T 2e4, 6, 7; Guinea, Liberia, Ivory
Coast, Ghana, Cameroon, Bioko, Sa ~o Tome , DR Congo, Rwanda,
Burundi, Ethiopia, Zambia, Malawi, Mozambique, Zimbabwe, South
Africa, Madagascar, Comoro Is., Reunion. At Kilimanjaro scattered,
nearly exlusively on the southern slope.
5.1.10. Lycopodiaceae
Terrestrial or epiphytic, erect to pendulous herbs or climbers
(Øllgaard, 1990).
5.1.10.1. Spore morphology. Spores trilete; heteropolar-radially
symmetric; sclerine fossulate; laesura 2/3 length or extending
almost to equator; amb subangular-convex; perine absent (Roubik
and Moreno, 1991).
Huperzia dacrydioides (Baker) Pic. Serm (Plate 2, 24e25).
Shape: triangular ~43 mm, the lobes usually truncate, and the
adjacent sides slightly depressed.
Aperture: trilete, laesura 2/3 to nearly equal the radius.
Surface: mostly foveolate, also fossulate.
Structure: exospore nearly plain on proximal face, apertural fold
slightly protruding, distal face usually pitted; perispore not seen.
Habitus: pendulous epiphyte or less often on rocks, rarely (not
in East Africa) terrestrial: stems 20e180 cm long.
Habitat: evergreen intermediate and montane forest, Ocotoea-
Podocarpus etc., riverine forest, forest patches and woodland, (900-
) 1500e2700 m. At Kilimanjaro epiphytic in moist lower-upper
montane Ocotea, Podocarpus and riverine forest, from 1400 to
2800 m.
Distribution: U 2, K 2e6; T 2, 3, 6e8; Sierra Leone, Guinea,
Liberia, Nigeria, Cameroon, Bioko, Sa ~o Tome, DR Congo, Burundi,
Sudan, Ethiopia, Malawi, Mozambique, Zimbabwe, South Africa,
Comoro Is. At Kilimanjaro rare, exlusively on the southern slope.
Huperzia ophioglossoides (Lam.) Rothm (Plate 2, 26e27).
Shape: slightly convex triangular ~42 mm.
Aperture: trilete, laesura 2/3 to nearly equal the radius.
Surface: plain proximal surfaces above and foveolate distal face
below.
Structure: exospore nearly plain on proximal face, apertural fold
slightly protruding, distal face usually pitted; perispore not seen.
Habitus: pendulous epiphyte or sometimes on rocks; stems (10-
) 30e150 cm long, dichotomously branched.
Habitat: upland evergreen forest especially Podocarpus,
Schefflera, Erica, bamboo etc. mist forest of Newtonia, Macaranga etc
(1600-) 1800e3250 m. At Kilimanjaro epiphytic in moist lower-
upper montane Ocotea, Podocarpus and riverine forest, from 1800
to 3000 m.
Distribution: U 2, 3; K 3, 4, 7; T 2, 3, 5e7; Bioko, Cameroon, DR
Congo, Rwanda, Burundi, Sudan, Ethiopia, Malawi, Mozambique,
Zimbabwe, South Africa, Madagascar, Re union, Mauritius and
Comoro Is. At Kilimanjaro scattered, exlusively on the southern
slope.
Huperzia saururus (Lam.) Trevisan (Plate 2, 28e29).
Shape: triangular ~50 mm, the lobes usually truncate.
Aperture: trilete, laesura 2/3 to nearly equal the radius.
Surface: foveolate.
Structure: exospore nearly plain on proximal face, apertural fold
slightly protruding, distal face usually pitted; perispore not seen.
Habitus: terrestrial or sometimes on rocks; horizontal stems
compactly branched. Aerial stems somewhat fleshy, mostly un-
branched, 9e60 cm tall.
Habitat: near streams and pools, Sphagnum swamps, boggy
moorland, seepage zones in rocks and exposed grassy crater rims,
Hagenia forest, 2200e4400 m. At Kilimanjaro in upper montane
Erica excelsa forest, subalpine Erica trimera heathland from 3100 to
3700 m.
Distribution: U 2, 3; K 3, 4; T 2, 6, 7; Cameroon, DR Congo,
Malawi, Zimbabwe, South Africa, Comoros, Re union, Mauritius,
Kerguelen Is., Tristan da Cunha and St. Helena, South America. At
Kilimanjaro very rare.
Lycopodium clavatum L (Plate 2, 30e32).
Shape: convex triangular ~33 mm (without reticulum).
Aperture: trilete, laesura 3/4 to nearly equal the radius.
Surface: reticulate with high (~5 mm) but thin muri.
Structure: exospore forming reticulate surfaces, the apertural
fold clearly projecting; perispore thin, usually of one layer.
Habitus: terrestrial with main stems creeping up to 1.5 m (but
sometimes ± lacking), producing dichotomously branched erect
stems 20e60 (80) cm tall at intervals of 5e10 cm.
Habitat: in both dry and wet habitats, bare earth, steep slopes
with gritty soil, roadside banks, grassland, moist montane forest
and rain forest, bamboo communities and extending into erica-
ceous heath e.g. Podocarpus, Ercia, Arundinaria, rocky places,
brackens scrub, swamps, in places can be dominant, (1400-)
1500e3050 m. At Kilimanjaro bimodal distribution avoiding the
dense and shady middle montane forest zone: from 1600 to 1900
and from 2400 to 3300 m. In the lower part of the forest belt a
pioneer species in open conditions on eroded soils, landslides,
recent road embankments and forest edges. In the upper part in
light uppermontane Erica excelsa forest and subalpine Festuca
obturbans tussock grasslands.
Distribution: U 2, 3; K 3e5, 7; T 2, 3, 6, 7; Nigeria, Cameroon,
Bioko, Sa~o Tome , DR Congo, Rwanda, Burundi, S Sudan, Ethiopia,
Zambia, Malawi, Mozambique, Zimbabwe, South Africa,
Madagascar, Mascarene and Comoro Is., ± cosmopolitan. At Kili-
manjaro scattered.
5.1.11. Marattiaceae
Terrestrial ferns. Marattiaceous ferns occur throughout the
tropics and subtropics (Camus, 1990).
5.1.11.1. Spore morphology. Spores monolete; heteropolar-bilateral;
sclerine echinulate; laesura 2/3 length, elliptical-acute; laesura
with thin margo; spinulae conspicuous; amb circular to elliptical
(eq) to planar convex (p); perine absent (Roubik and Moreno, 1991).
Marattia fraxinea Sm (Plate 2, 33e34).
Shape: ellipsoidal ~31  35 mm.
Aperture: monolete, linear, 1/2 to 2/3 the length, often obscure.
 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
Surface: coarsely rugate.
Structure: exospore of three layers, the contours largely formed
by the exospore; perispore conforms to the exospore and is locally
divided into two strata.
Habitus: rhizome forming an erect caudex, up to 40 cm high and
30 cm across. Fronds tufted, arching, (1-) 2.5e4 m tall.
Habitat: evergreen forest (particularly primary), montane forest,
often in swampy places, sometimes riverine, 900e2550 m. At
Kilimanjaro mainly in submontane-lower montane Mitragyna,
Newtonia and Afrocrania riverine and gorge forests, together with
Dryathyrium boryanum and Didymochlaena, with which it shares
habitat and distribution, from 1200 to 2300 m.
Distribution: U 2, 4; K 4, 5, 7; T 1e4, 6e8; widespread in Africa
from Guinea and Sierra Leone to Ethiopia and Angola, South Africa,
Madagascar, Mascarene Is. At Kilimanjaro scattered and only on the
wet southern slope.
5.1.12. Ophioglossaceae
Mainly terrestrial herbs, some epiphytic usually small and fle-
shy, lacking sclerenchyma. Except for Helminthostachys, the genera
are nearly cosmopolitan (Wagner, 1990).
5.1.12.1. Spore morphology. spore trilete; heteropolar-radially
symmetric; sclerine striate-verrucate, striate-scabrate; laesura 1/
2 of the radius with complete ‘Y’ split mark; laesura with con-
spicuous margo; amb circular; perine present, irregular, bacculate-
perforate (Roubik and Moreno, 1991).
Ophioglossum vulgatum subsp. kilimandscharicum (Hieron.) J.E.
Burrows (Plate 2, 35e37) SYN Ophioglossum petiolatum Hook.
Shape: spheroidal, triangular convex ~48 mm.
Aperture: trilete, rarely monolete, the laesura 1/2 to 2/3 the
radius.
Surface: irregular rugae more or less fused in a reticulum,
somewhat denser on the proximal face, with many small granules
on the surface that are echinate in profile.
Structure: exospore of three main layers: the inner compact, the
middle with many cavities and foliations, and the outer largest,
compact stratum; a thin perispore of fine fibrillar material is easily
detached.
Habitus: perennial herb with deciduous aerial parts. Rhizome
fusiform to linear, (6-) 10e18 (30) mm long, 2e4 mm wide; roots
few, fleshy, horizontal, sometimes descending, proliferous.
Habitat: in montane grassland or scrub, particularly among
large grass tussocks (e.g. Miscanthus sp.) in marshy areas;
Plate 3. POLYPODIACEAE: 1 Loxogramme lanceolata, 2e4 Lepisorus excavatus, PTERIDACEAE: 5e8 Pteris dentata, SELAGINELLACEAE: 9e11 Selaginella goudotiana var. abyssinica,
THELYPTERIDACEAE: 12e14 Amauropelta bergiana, 15e17 Christella dentata, WOODSIACEAE (Cystopteridaceae): 18e20 Cystopteris fragilis ssp. diaphana.
321
2200e2800 m. At Kilimanjaro in humid, anthropogenic
submontane-lower montane meadows and grasslands, inside the
forest belt on disturbed ground, e.g. landslides and roadsides, from
1400 to 2100 m.
Distribution: U 2, K 5; T 2; DR Congo and South Africa. At Kili-
manjaro very rare.
5.1.13. Polypodiaceae
Usually terrestrial or epiphytic, sometimes epilithic, usually
small to medium-sized, sometimes large ferns with short-to long-
creeping stem bearing basifixed (Hennipman et al., 1990).
5.1.13.1. Spore morphology. Spores monolete, heteropolar-bilateral
and trilete, heteropolar-radially symmetric; sclerine psilate, sca-
brate, granulate, verrucate, fossulate; laesura always narrow (trilete
type) and elliptical-acute (monolete type), 2/3 of radius; margo
wide and conspicuous; amb subangular-convex (eq) and convex-
planar to biconvex (p) in trilete spores, convex-planar and
concave-convex in monolete spores; perine present, conspicuous in
50% of the species; psilate, scabrate, reticulate, granulate, echinu-
late, bacculate (Roubik and Moreno, 1991).
Loxogramme lanceolata (Plate 2, 38e39 & Plate 3, 1) SYN
L. abyssinica (Baker) M.G. Price.
Shape: ellipsoidal ~64  86 mm.
Aperture: trilete sometimes with unequal laesura, 1/2 to 3/4 the
radius, or monolete, 1/4 to 2/3 the length.
Surface: papillate to somewhat rugate.
Structure: exospore forms the surface contours; perispore usu-
ally thin, laminate, some parts thickened.
Habitus: epiphytic or sometimes on rocks. Rhizome widely
creeping.
Habitat: evergreen forest, mixed Juniperus-Podocarpus forest,
occasionally river banks and rocky places but mostly epiphytic,
900e2900 m. At Kilimanjaro mainly epiphytic in moist lower-
upper montane Ocotea, Podocarpus and riverine forest together
with Pleopeltis excavata and P. macrocarpa, in lower elevations on
boulders in stream bed inside riverine forest, from 980 to 2900 m.
Distribution: U 1e4; K 1, 3e7; T 2e4, 7, 8; widespread
throughout Africa to South Africa, also Comoro Is. and Madagascar.
At Kilimanjaro widespread.
Lepisorus excavatus (Bory ex Willd.) Ching (Plate 3, 2e4) SYN
Pleopeltis excavata (Bory ex Willd.) Sledge.
Shape: ellipsoidal ~56  78 mm.
Aperture: monolete, 2/3 to 3/4 the length.
322 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
Surface: verrucate, usually low and thick verrucae.
Structure: very thick exospore forms the surface contours.
Habitus: epiphytic or often on rocks. Rhizome creeping with
fronds borne 4e12 mm apart.
Habitat: upland and montane evergreen forest, often riverine,
particulary Olea-bamboo, Olea-Diospyros, Trichocladus-Cassipourea-
Aphloia, Parinari, Mitragyna etc., Ocotea-Podocarpus valley forest,
also Brachystegia woodland and tree-heath montane zone,
1150e3200 m. At Kilimanjaro epiphytic mainly in moist lower-
upper montane Ocotea, Podocarpus, Erica excelsa and riverine for-
est together with Loxogramme abyssinica and P. macrocarpa, also in
coffee-banana plantations (Chagga homegardens), from 1300 to
3500 m.
Distribution: U 1e3: K 1e7; T 2e8; Sierra Leone to Angola,
Ethiopia, Sudan, DR Congo-Kinshasa, Rwamda, Zambia, Malawi,
Mozambique, Zimbabwe, South Africa, Madagascar, Mauritius,
Reunion, Comoro Is. At Kilimanjaro widespread.
5.1.14. Pteridaceae
Terrestrial or rupestral, or (Ceratopteris) aquatic ferns of mostly
small size, exceptionally leaves are to 6 m long. Pteridaceae are
essentially worldwide in distribution, although centred in the
Tropics (Tryon et al., 1990).
5.1.14.1. Spore morphology. Spores trilete, rarely alete (Tryon et al.,
1990).
Pteris dentata Forrsk (Plate 3, 5e8).
Shape: triangular ~38 mm.
Aperture: trilete, the open laesura 2/3 to ¾ the radius, rarely
monolete.
Surface: echinate and verrucate, sometimes forming a reticuloid
pattern.
Structure: exospore of two layers, the outer forming the main
contours with numerous canals, sometimes large in the apertural
region, perispore of two layers conforming to the exospore con-
tours, the lower thin and finely granular, the outer rougher, forming
rodlets in some species and sometimes enclosing globules.
Habitus: terrestrial or rarely epiphytic (truly?); rhizome erect or
creeping. Fronds tufted, 0.15e2.1 m tall.
Habitat: grassland, upland bamboo forest, montane rain forest,
often by stream sides, also old rubber plantations, bare slopes of
landslips and on trunks of oil palms, 1000e2700 (2950) m. At
Kilimanjaro mainly in moist submontane-lower montane Mitra-
gyna, Newtonia and Afrocrania riverine and gorge forests, in lower
montane Cassipourea forest, also on humid shaded embankments
of irrigation funnels and road ditches in the Chagga homegardens
and here growing together with A. hispidulum, A. raddianum and
A. incisum, from 960 to 2600 m.
Distribution: U 2e4; K 1, 3e7; T 1e4, 6e8; Cape Verde Is., Ghana,
Bioko, E. DR Congo, Rwanda, Burundi, Ethiopia, Sudan, Angola,
Zambia, Malawi, Zimbabwe, South Africa, Madagascar, Mascarene
Is., Ascension I., St. Helena, Greece, Crete, Arabia. At Kilimanjaro
widespread.
5.1.15. Selaginellaceae
Terrestrial, or very occasionally epiphytic, usually perennial
plants, of varied habitus.
A family of a single genus, with some 750 species mainly in the
tropical zone of the world, with a few species reaching the arctic-
alpine zones in both hemispheres (Jermy, 1990).
5.1.15.1. Spore morphology. Spores trilete, heteropolar radially
symmetric, sclerine echinulate, bacculate, sexine elements con-
spicuous, laesura narrow, 2/3 of length, always distinct ‘Y’ mark at
the ends, amb subangular-convex to almost circular, perine absent
(Roubik and Moreno, 1991).
Selaginella goudotiana var. abyssinica (Spring) Bizzarri (Plate 3,
9e11).
Shape: slightly convex triangular ~36 mm.
Aperture: trilete, the open laesura 3=4 to nearly equal the radius,
with thin margo.
Surface: coarsly rugulate/verrucate.
Structure: exospore plain, verrucate, or spinulose, usually
overlaid by either perispore or paraexospore, or sometimes both
may be lacking.
Habitus: plant erect or suberect with ascending branches,
3e40 cm long or tall. Sometimes in dense stands.
Habitat: on rocks, near streams, riverbanks, waterfalls and floor
layer in evergreen forest; also damp grassy slopes; 750e2450 m. At
Kilimanjaro mainly in moist submontane-lower montane Mitra-
gyna, Newtonia and Afrocrania riverine and gorge forests, on wet
(watersprayed) shaded rocks in riverine forests together with
Adiantum capillus-veneris and Pilea rivularis, also on humid shaded
embankments of irrigation funnels and road ditches in the Chagga
homegardens and here growing together with A. hispidulum, A.
raddianum and A. incisum, from 1000 to 2500 m.
Distribution: U 3, K 2e5, 6, 7; T 2e8; At Kilimanjaro scattered,
only on the southern slope.
5.1.16. Thelypteridaceae
Terrestrial (rarely epiphytic) ferns with branched or unbranched
stems, these long- or short-creeping, suberect, or erect and trunk-
like. Thelypteridaceae are one of the largest families of ferns,
comprising nearly 1000 species mostly in tropical and subtropical
regions.
5.1.16.1. Spore morphology. Spores nearly always bilateral with a
monolete scar (tetrahedral with trilete scar in Trigonospora), peri-
spore variously ornamented (reticulate, winged, spinulose) (Smith,
1990).
Amauropelta bergiana (Schldl.) Holttum (Plate 3, 12e14).
Shape: ellipsoidal 33  42 mm.
Aperture: monolete, 2/3 to 3/4 the length with thin margo.
Suface: (loosely) echinate, papillate.
Structure: exospore gemmulate, perispore two layered, the
outer usually cavate or with large surface discontinuities.
Habitus: rhizome erect. Fronds tufted, up to 1e1.2 m tall and
25 cm wide.
Habitat: wet evergreen forest, bamboo-Podocarpus, usually in
boggy areas by streams, rock caves in Erica forest, 1200e2500
(2600?) m. At Kilimanjaro in most humid forest types from the
colline to the upper montane zone, also in coffee-banana planta-
tions (Chagga homegardens), from 800 to 3200 m.
Distribution: K 4, 5, 7; T 2e4, 6, 7; Cameroon, Bioko, Rwanda,
Burundi, Sudan, Ethiopia, Zambia, Malawi, Zimbabwe, South Africa,
Mascarene Is. At Kilimanjaro widespread.
Christella dentata (Forssk.) Brownsey & Jermy (Plate 3, 15e17).
Shape: ellipsoidal ~28  38 mm.
Aperture: monolete, 2/3 to 3/4 the length.
Surface: coarse folds, cristate.
Structure: exospore with somewhat gemmulate outer layer,
perispore heterogeneous, the outer layer more or less cavate.
Habitus: rhizome shortly to distinctly creeping, ± 7 mm in
diameter. Fronds closely spaced, 0.4e1.5 (2) m tall.
Habitat: Evergreen forest, often riverine, ditch-sides in swampy
areas, valley bushland and thicket; (0?-) 45e2100 m. At Kilimanjaro
in moist submontane-lower montane Mitragyna, Newtonia and
Afrocrania riverine and gorge forests, however mainly as an apo-
phyte in anthropogenic vegetation (humid shaded embankments of
irrigation funnels and road ditches in the Chagga homegardens),
 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
from 700 to 2100 m.
Distribution: U 2e4; K 2e5, 7; T 1e4, 6, 8; throughout the Old
World tropics and subtropics, just reaches into SW Spain; intro-
duced in some parts of tropical America. At Kilimanjaro wide-
spread, but mainy on the southern and eastern slope.
5.1.17. Woodsiaceae
Terrestrial with erect or less often creeping rhizome, sometimes
forming a short caudex. A cosmopolitan family containing about 15
genera and 500 species.
5.1.17.1. Spore morphology. Spore monolete with perispore
(Verdcourt, 2003).
Cystopteris fragilis ssp. diaphana (Bory) Litard (Plate 3, 18e20).
SYN Cystopteris fragilis (L.) Bernh. Ssp B. Verdc. (Cystopteridaceae)
(in Kubitzki system: Drypteridaceae Athyrioideae).
Shape: ellipsoidal ~30  46 mm.
Aperture: monolete, 2/3 to 3/4 the length.
Surface: coarsly echinate with the echinae projecting from a
rugate surface.
Structure: exospore plain, contours formed by granulate
perispore.
Habitus: rhizome shortly creeping or ± erect. Fronds ± tufted or
closely spaced.
Habitat: mostly on clamp rocks, rock faces and cliffs, often near
rivers, chiefly evergreen forests, e.g. Hagenia, Podocarpus, Albizia-
Macaranga-Fagara-Syzygium-Croton or Xymalos-Prunus-Maesa
Afrocrania; also on moist loamy banks and alpine zone;
1400e3800 m. At Kilimanjaro mainly (on rocks) in lower-upper
montane riverine forests, from 1500 to 3700 m.
Distribution: U 2, 3; K 1, 3e5; T 2, 7; Cameroon Mts, DR Congo,
Sudan, Ethiopia, Natal, Mediterranean part of Europe, Cape Verde
Is., Canary Is. At Kilimanjaro scattered.
5.2. Pollen types
5.2.1. Acanthaceae
The family is comprised of shrubs, herbs and trees (rarely, but
including a few mangroves). It consists of about 250 genera and
2500 species. The distribution is Holarctic, Palaeotropical,
Neotropical, Cape, and Australian, in temperate to tropical climate.
The family is centred on Indomalaysia, Africa, Brazil and central
America (Watson and Dallwitz, 1992 onwards).
5.2.1.1. Pollination. Entomophilous, mechanism conspicuously
specialized or unspecialized (Watson and Dallwitz, 1992 onwards).
5.2.1.2. Pollen morphology. The family is distinctly eurypalynous
with radiosymmetric, or less frequently bilateral, 3-8-colpate, 3-5-
colporate, 2-4-porate, periporate or spiraperturate pollen grains.
Colporate pollen grains are sometimes provided with pseudocolpi;
peroblate-peroblate (Erdtman, 1952).
Asystasia gangetica (L.) T. Anderson (Plate 4, 1e2).
Shape: prolate, rectangular ~35e40  50e65 mm, triangular
~54 mm.
Aperture: 3-colporate, pore round with lolongate annulus, colpi
long (full) with thick margo.
Exine: coarsely reticulate, areolate, heterobrochate, with a sin-
gle row of bacule, sexine thicker than nexine, exine thicker at the
equatorial area.
Habitus: perennial or suffrutescent herb with irregular root-
stock, occasionally rooting from lower nodes. Stems creeping-
ascending or straggling, to 2 (3) m long.
Habitat: forest margins and clearings and a wide range of variety
of woodland, wodded grassland and bushland, sandy sea shores,
323
roadsides, ditches, plantations, often in disturbed or secondary
vegetation; 0e1700 m. At Kilimanjaro in wet and dry grasslands
and anthropogenic meadows, in ruderal vegetation (Chagga
homegardens, roadsides, fallow arable fields and waste places)
from the savanna to the lower montane zone, rarly in forests, from
950 to 1680 m.
Distribution: U 1e4; K 1e7; T 1e8; Z; P; throughout tropical
Africa and South Africa, Madagascar, Mascarene Is., tropical Arabia,
introduced into Malaysia. At Kilimanjaro scattered.
Pollination: mainly two bee genera: Amegilla and Lasioglossum
(Kato et al., 2008).
Blepharis maderaspatensis B.Heyne ex Roth (Plate 4, 3e5).
Shape: prolate ~24  44 mm, circular to triangular.
Apertures: 3-colpate, colpi long (4/5) and narrow.
Exine: perforate (small simple, evenly spaced perforations) to a
rough hexagonal reticulum, exine semitectate with simple collu-
mellae, a foot layer and thick endexine which is fractioned at the
colpi (thickness: endexine: 1e2 mm, ectexine: 0.5e1 mm).
Habitus: suberect, procumbent or scrambling perennial e rarely
annual e herb, rarely a suberect or trailing subshrub. Stems up to
2 m long, sometimes rooting at nodes.
Habitat: wide range of grassland, bushland, woodland and forest
habitats, also commonly in disturbed and secondary vegetation and
ruderal; 0e1850 m. At Kilimanjaro in dry savanna forests and
grasslands and dry submontane anthropogenic grasslands, in
ruderal vegetation (Chagga homegardens, roadsides, fallow arable
fields and waste places), from 820 to 1580 m.
Distribution: U 1e3; K 1e7; T 1e8; Z; P; widespread in tropical
Africa and through Arabia to India and Thailand. At Kilimanjaro
rare.
Pollination: insects. No further information known.
Isoglossa laxa Oliv (Plate 4, 6e9).
Shape: bilateral, circular/ellipsoidal/rectangular ~34  40 mm.
Apertures: 2-porate with a more or less pronounced marginal
‘girdle’, pore Ø ~4 mm.
Exine: crassinexinous, reticulate, rambiculate with crassinex-
inous aperture margins.
Habitus: perennial herb or subshrub, 0.3e2 m tall.
Habitat: montane forest understorey, clearences and secondary
growth, often dominant. At Kilimanjaro widespread in submontane
to middle montane forests (riverine and gorge forests, Croton-Cal-
odendrum, Cassipourea, Ocotea forests), from 1300 to 2740 m.
Distribution: K 6; T 2; not known elsewhere. At Kilimanjaro
inside the forest belt widespread.
Pollination: insects. No further information available. Apis mel-
lifera adansonii has been observed pollinating the African Isoglossa
woodii (Griffiths et al., 2010).
Justicia asystasioides (Lindau) M.E.Steiner (Plate 4, 10e12).
Shape: radiosymmetric, prolate ~36  49 mm, circular.
Apertures: heterocolporate, 2-colpori-4-pseudocolpate, pori
circular Ø ~3 mm with well-defined costa, colpi with pori wider
around pori, colpi without pori evenly narrow colpi length 5/6.
Exine: tectate, tectum perforate, supra-reticulate in the pe-
ripheral area, homobrochate, muri simply-collumellate.
Habitus: erect or straggling herb or shrub to 3 (4) m tall.
Habitat: wet evergreen intermediate and montane forest; (759-)
950e2000 (? 2900) m. In the North Pare Mountains and the West
Usambaras in Ocotea forest.
Distribution: T 3, 6e8; Malawi, Mozambique.
Pollination: insects. Species of stingless bees (Meliponinae) and
skipper (Hesperiidae) (Do € ll et al., 2007), hummingbirds and honey
bees (Quintana-Va squez, M. de los Angeles, 2007) as well as
Diptera, Hymenoptera and Lepidoptera (McMullen, 1994) have
been observed pollinating Central and South American Justicia
species.
324 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386

Plate 4. ACANTHACEAE: 1e2 Asystasia gangetica, 3e5 Blepharis maderaspatensis, 6e9 Isoglossa laxa, 10e12 Justicia asystasioides, 13e17 Justicia diclipteroides, 18e20 Thunbergia alata,
AMARANTHACEAE: 21e23 Achyranthes aspera, 24e26 Amaranthus hybridus, 27e28 Celosia schweinfurthiana, ANACARDIACEAE: 29e31 Lannea triphylla, 32e35 Ozoroa insignis,
36e39 Rhus vulgaris, ANTHERICACEAE: 40e41 Chlorophytum rhizopendulum, 42e44 Chlorophytum viridescens; APOCYNACEAE: 45e47 Carissa edulis, 48e49 Rauvolfia caffra, 50e52
Saba comorensis, 53 Tabernaemontana stapfiana.

Justicia diclipteroides Lindau (Plate 4, 13e17).
Shape: radiosymmetric, prolate ~25  40 mm, circular.
Apertures: dicolporate, pori elliptic broad ~2  3 mm with well-
defined costa, colpi long (4/5) and narrow.
Exine: tectate, collumellate, very thick collumella, thickend
around equator, hence exine thinner on poles, reticulate,
homobrochate.
Habitus: perennial or shrubby herb with several suberect,
ascending, decumbent or scandent stems from erect to creeping
rhizome, to 60 cm long in erect plants, to 2 (3) m long in scandent
ones.
Habitat: montane evergreen forest, particularly in glades and on
margins, montane grassland and bushland, lowland and interme-
diate bushland, rocky hills, riverine forest; (250-) 550e2600 m. At
Kilimanjaro in dry forests and riverine forests of the savanna,
submontane Croton-Calodendrum forests, lower-middle montane
Cassipourea forests and upper montane Juniperus forests, avoiding
moist forest types of the southern slope, from 1030 to 2900 m.
Distribution: U 1; K 1e4, 6, 7; T 1e5, 7; Ethiopia, DR CONGO,
Rwanda, Burundi. Inside the forest belt Kilimanjaro scattered,
mainly on the drier western and northern slope.
Pollination: See J. asystasioides.
Thunbergia alata Sims (Plate 4, 18e20).
Shape: circular, spheroidal ~64 mm.
Apertures: spiraperturate.
Exine: psilate, scabrate.
Habitus: annual or perennial trailing or twining herb. Stems to
5 m long.
Habitat: wide range of habitats from wet forest (usually on
margins) to dry Acacia-Commiphora bushland, often in disturbed
places and secondary vegetation; near sea level to 2700 m. At
Kilimanjaro in wet and dry anthropogenic grasslands and
meadows, in ruderal vegetation (Chagga homegardens, roadsides,
fallow arable fields and waste places) from the colline to the lower
montane zone, in forest clearings, rarly in forests, from 970 to
2470 m.
 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
Distribution: U 1e4; K 1e7; T 1e8; Z; P; throughout tropical
Africa, especially common in the East, Madagascar, India, SE Asia,
introduced in tropical America. At Kilimanjaro scattered.
Pollination: pollinators are unknown. In South Africa this spe-
cies is pollinated by bees and the flowers reflect ultra violet light in
a pattern that is only visible to insects (Schmidt-Lebuhn et al.,
2007).
5.2.2. Amaranthaceae
Annual or perennial herbs, subshrubs or shrubs, rarely lianes or
trees. A mainly tropical family (but found in all continents) of 69
genera and over 1000 species (Townsend, 1993).
5.2.2.1. Pollination. Insect pollinated, except for Amaranthus (wind
pollinated) (Townsend, 1993).
5.2.2.2. Pollen morphology. Erdtman (1952) demonstrated two
major pollen types in this family, termed Amaranthus type and
Gomphrena type. Pollen grain in the Amaranthaceae are always
periporate, the Amaranthus type having a microechinate and tubi-
liferous/punctate ectexine. The Gomphrena type has a reticulate
ectexine with the pore deeple recessed.
Achyranthes aspera L (Plate 4, 21e23).
Shape: spheroidal ~ 22 mm.
Aperture: periporate, circular pori with thin annulus, pori
2.0e4.0 mm in Ø with a ±irregular borderline to the tectum.
Exine: tectate, perforate, columellate, scabrate, psilate.
Habitus: perennial herb e sometimes woody and somewhat
suffrutescent e, 0.2e2 m, stiffly erect to subscandent or straggling.
Habitat: practically all habitats from coastal scrub to dry cliff
tops to mist forest; common as a weed of cultivation or disturbed
ground, in open grassland, hardpan patches between rocks, along
forest trails and edges, in seasonal swamps and dried-up water-
courses, etc.; grows on a wide range of soil types from red sandy
ground to black cotton soil or granite mountain tops; 0e3000 m. At
Kilimanjaro in dry forests and riverine forests of the savanna,
submontane Croton-Calodendrum forests, lower-middle montane
Cassipourea forests and upper montane Juniperus forests, avoiding
moist forest types of the southern slope, outside forests mainly in
ruderal vegetation (Chagga homegardens, roadsides, fallow arable
fields and waste places), from 780 to 3010 m.
Distribution: found throughout the world in tropical and
warmer regions generally. Some forms may have originated in
more restricted areas but these have now become sufficiently
widely dispersed to make this a matter of conjecture. At Kili-
manjaro widespread, inside the forest belt mainly on the drier
western and northern slope.
Pollination: mainly bee pollinated (Wubie et al., 2014).
Amaranthus hybridus L. agg (Plate 4, 24e26).
Shape: spheroidal, circular ~25 mm.
Aperture: periporate (30e65 pori), circular pori, no annulus
visible, pore 1.0e2.0 mm in diameter, sunken and sharply set off
against the tectum.
Exine: psilate to scabrate, tectate perforate with very thin col-
lumellae and with numerous evenly distributed microspines.
Habitus: annual herb, erect or less commonly ascending, up to
±2 (3) m in cultivated forms, but much less in spontaneous plants.
Habitat: commonly as a weed of current or abandoned cultiva-
tion or waste ground, also along roadsides, river margins and forest
edges; 900e2210 m. At Kilimanjaro only synanthrop in ruderal
vegetation (Chagga homegardens, roadsides, fallow arable fields
and waste places), from 800 to 1800 m.
Distribution: U 2, 4; K 1, 3e6; T 1e8; Z; throughout the tropical
and subtropical regions of the world, also occurring as a casual in
temperate regions, e.g. a common wool adventive in Europe,
325
originated in America. At Kilimanjaro below the forest belt
widespread.
Pollination: wind (Arntz et al., 1998).
Celosia schweinfurthiana Schinz (Plate 4, 27e28).
Shape: spheroidal, circular ~24 mm.
Apertures: periporate, pori Ø 4e5 mm with a ±irregular
borderline to the tectum, mesoporia flat.
Exine: tectate, perforate with few evenly distributed micro-
spines, psilate to scabrate.
Habitus: perennial herb, often suffrutescent, varying consider-
ably in habitus from a prostrate plant rooting at the nodes to an
erect herb 15 cm tall or a climber rambling to 5 m.
Habitat: as ground cover along forest rides, margins and
clearing, especially near water or scrambling in thicker forest,
otherwise in roadside or coastal bushland, or as weed of cultiva-
tion; 3e1550 m. At Kilimanjaro mainly in ruderal vegetation
(Chagga homegardens, roadsides, fallow arable fields and waste
places), rarely in colline-lower montane forests (riverine, Croton-
Calodendrum, Cassipourea forest), from 820 to 1780 m.
Distribution: U 2e4; K 1, 3, 5, 7; T 1e3, 5e7; Sudan, Ethiopia,
Zaire, Angola. At Kilimanjaro rare, inside the forest belt mainly on
the drier western and northern slope.
Pollination: insects. No further details known.
5.2.3. Anacardiaceae
Trees, shrubs, rarely subshrubs, lianas. Approximately 81 genera
and 800 species in dry to moist, mostly lowland habitats in the
tropics and subtropics worldwide, but also extending into the
temperate zone (Pell et al., 2011).
Pollination: Anacardiaceae are primarily entomophilous, but
some exceptions are found (Anacardium: butterflies and moths,
some Mangifera). A few genera are wind-pollinated (Amphiptery-
gium, Campylopetalum, Dobinea, Orthopterygium, Pistacia (Pell et al.,
2011).
Pollen morphology: pollen grains are generally 3-colporate,
spheroidal, the colpi are long and narrow. The reticulation is coarse,
with high upstanding ridges enclosing large polygonal lacunae. The
sexine can be finely grano-rugulate, striate-perforate or reticulate.
The pore shape varies from spherical to oblong with a smooth,
ragged or psilate surface (Heimsch, 1940; Erdtman, 1952;
Marticorena, 1968; Baksi, 1976; Ibe and Leis, 1979; Olivera et al.,
1998).
Lannea triphylla (A. Rich.) Engl (Plate 4, 29e31).
Shape: ±spheroidal ~26  30 mm, circular.
Apertures: 3-colporate, apertures weakly defined, pori tall
elliptic, colpi length 2/3, colpal membrane granulate.
Exine: tectate, finely striate.
Habitus: deciduous shrub or small tree, 2e5 m high (Thulin,
1999).
Habitat: in wooded grassland and deciduous bushland, often on
stony hills; 340e1400 m. At Kilimanjaro in savanna woodland, from
900 to 1000 m.
Distribution: U 1, K 1e4, 6, 7; T 3; Ethiopia, Somalia, Arabia. At
Kilimanjaro rare, in the savanna zone scattered.
Pollination: unknown; probably insect pollinated like in most
other Anacardiaceae since not observed otherwise.
Ozoroa insignis Delile (Plate 4, 32e35).
Shape: rhombic tall ~30  35 mm, circular ~38 mm.
Apertures: 3-colporate, pori rectangular broad ~3  10 mm,
mesoporia slightly vaulted, colpi 4/5 in length and narrow.
Exine: tectate, striato-microreticulate.
Habitus: a much branched tree or shrub 3e15 m high.
Habitat: wooded grassland, often on scarps and rocky hills,
extending marginally to drier forests and higher rainfall deciduous
woodland and bushland; 5e2200 m. At Kilimanjaro in savanna
326 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
woodland and dry forest, from 890 to 1200 m.
Distribution: U 1e4; K 1e7; T 1e8; Zaire, Burundi, Rwanda, S.
Sudan, S. Ethiopia, Malawi, Mozambique, Zimbabwe, Zambia,
Angola, and South Africa. At Kilimanjaro rare, in the savanna zone
scattered.
Pollination: probably insects like other Ozoroa species.
Rhus vulgaris Meikle (Plate 4, 36e39).
Shape: subprolate ~18  25 mm, trilobate ~18 mm.
Apertures: 3-colporate, colpi long (5/6) and narrow, seemingly
broad rectangular (~7  1.5 mm) pori but they are actually colpi
transversales with costae. Costae not to the end of colpi.
Exine: tectate, columellate, striate.
Habitus: shrub or tree up to 10 m high.
Habitat: upland evergreen bushland, forest, edges, on hill slopes
and valleys, on lake shores and along river banks, and wooded
grassland; 800e2700 m. At Kilimanjaro in savanna woodland,
submontane Croton-Calodendrum forest, secondary bushland,
from 1000 to 1750 m.
Distribution: U 1e4; K 1e6; T 1e8; Cameroon to Ethiopia, also in
Mozambique, Malawi, Zimbabwe, and Zambia. At Kilimanjaro
scattered.
Pollination: flowers of Rhus spp. are visited by several species of
bees, flies and other insects, suggesting that many generalist pol-
linators influence the reproductive success of plants (Matsyuama
et al., 2009).
5.2.4. Anthericaceae
Perennial, terrestrial herbs from short rhizomes, rarely with a
woody stem (Chlorophytum suffructicosum). A family of worldwide
distribution of about 250 species in 22 genera distributed mainly in
Africa, Europe, Asia and the Americas, extending to N Australia
(Conran, 1998).
5.2.4.1. Pollination. Flowers of Anthericaceae are insect pollinated.
Kativu (1994) found that Zimbabwean Chlorophytum species are
self-compatible, although most taxa studied required active polli-
nation by various species of bees (Conran, 1998).
5.2.4.2. Pollen morphology. Pollen grains are sulcate. The sexine is
either thicker or thinner than the nexine. Exine sculpturing is
reticulate and homobrochate or heterobrochate, more or less
smooth (e.g. Chlorophytum, Echeandoa) or with raised sculpturing
(e.g. Anthericum, Paradisea) (Schulze, 1982; Diaz Lifante et al., 1990).
Chlorophytum rhizopendulum Bjorå & Hemp (Plate 4, 40e41).
Shape: prolate to tall rectangular ~16  28 mm, circular to
elliptic.
Apertures: monosulcate, narrow sulcus 4/5 to 5/5, often with
grainy and lacterated margins.
Exine: exine is very thin, psilate to slightly scabrate.
Habitus: perennial herb, rhizomes up to 1 m long, pendulous.
Habitat: grows on rocky walls of a gorge with tree cover within a
savanna area. At Kilimanjaro only on semi-shaded cliffs in savanna
riverine forests with Lecaniodiscus fraxinifolius, Albizia petersiana
and Trichilia emetica, from 900 to 1010 m.
Distribution: T2, endemic to Kilimanjaro, extremely rare.
Pollination: the genus Chorophytum is insect pollinated, by
various species of bees (Conran, 1998).
Chlorophytum viridescens Engl (Plate 4, 42e44).
Shape: (sub) prolate ~27  36 mm, circular to elliptic ~27 mm.
Apertures: monosulcate, sulcus (4/5) runs in the center of the
grain, tappering towards the end wider at equator.
Exine: exine is very thin, psilate to slightly scabrate.
Habitus: perennial herb. Plants up to 120 cm.
Habitat: rocky outcrop, on shallow black clayish soils;
500e2700 m. At Kilimanjaro in lower montane Ocotea and riverine
forest. Rarely in savanna grasslands, 1300e1400 m.
Distribution: U 1; K 3, 4, 6, 7; T 2, 3; Uganda (Acholi), Kenya
(Trans-Nzoia, Namuki, Teita), Tanzania (Masai, Mbulu, Handeni),
Cameroon. At Kilimanjaro rare.
Pollination: the genus Chorophytum is insect pollinated, by
various species of bees (Conran, 1998).
5.2.5. Apocynaceae
Trees (few, e.g. in Tabernaemontana, Dyera), or shrubs, or lianas
(mostly), or herbs (e.g. Vinca). Temperate (a few), or sub-tropical to
tropical (mainly). There are 1500 species in 164 genera (Watson
and Dallwitz, 1992 onwards).
Pollination: entomophily is predominant. There are some re-
ports on species of Apocynaceae pollinated by butterflies (e.g.
Waddington, 1976), flies and bees (e.g. Bawa et al., 1985), hawk-
moths (Haber, 1984), and hummingbirds (Linhart and Feinsinger,
1980).
Pollen morphology: pollen grains are usually 3-colporate or 2-3-
porate, more or less flattened to spherical. Grains are sometimes
irregularly pitted; the inner exine surface is often uneven. The exine
stratification is often obscure. The ± fused state of the sexinous
element is noteworthy (Erdtman, 1952).
Carissa edulis (Plate 4, 45e47) SYN C. spinarum
Shape: spheroidal to suboblate ~27  34 mm, circular to weakly
trilobate ~29 mm.
Apertures: 3-colporate, lolongate/tall elliptic pori, colpi (1/2)
narrow with costae.
Exine: psilate, scabrate, tectate, fissured.
Habitus: shrub 0.2e3 m, occasionally scrambling up to 20 m
high.
Habitat: miombo woodland, bushland, riverine forest or thicket,
upland forest especially rocky places; 0e2250 m. At Kilimanjaro in
grasslands, woodlands and secondary bushlands of the savanna
and submontane zone, from 1040 to 1750 m.
Distribution: U 1e4; K 1e7: T 1e7; widespread in tropical Africa,
tropical Asia and Australia. At Kilimanjaro scattered.
Pollination: insects; hawkmoths (Martins and Johnson, 2009).
Rauvolfia caffra Sond (Plate 4, 48e49).
Shape: suboblate to oblate, circular to slightly convex triangular
~49 mm.
Apertures: 3-colporate, colpi short (1/3) with margo, arcuate
with arci considerably thinner towards poles, pori lolongate with
thick annulus.
Exine: tectate, psilate, scabrate.
Habitus: shrub or tree 2e40 m high.
Habitat: riverine forest, swamp forest, moist forest;
450e1950 m. At Kilimanjaro mainly in riverine and gorge forests
(colline-lower montane), also frequently in Chagga homegardens,
from 800 to 1980 m.
Distribution: U 1e4; K 1, 3e7; T 1e8; widely distributed in
tropical Africa from Togo to Angola and from Sudan to South Africa.
At Kilimanjaro scattered.
Pollination: butterflies and other insects, which in turn attract
insect-eating birds that bring about pollination (Coates Palgrave
et al., 2002).
Saba comorensis (Bojer ex A. DC.) Pichon (Plate 4, 50e52).
Shape: spheroidal, circular ~34 mm.
Apertures: 1-pseudocolpato-2-colporate, pori are large, circular
Ø ~10 mm and annulate.
Note: this species can be provided with one, two or three ap-
ertures. In the first case the grains are provided with two pseudo-
colpi and one normal, porate colpus; in the second case they are 1-
pseudocolpato-2-colporate; in the third (and rare) case 3-colporate
(see also Erdtman, 1952). The second type is mainly found at
Kilimanjaro.
 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386 327

Exine: psilate; the exine stratification is rather obscure.
Habitus: liana up to 20 m long.
Habitat: moist forest, forest edges, secondary forest, thicket,
woodland in rocky sites; 0e2000 m. At Kilimanjaro in riverine and
gorge forests of the savanna and submontane zone, from 780 to
1220 m.
Distribution: U 1e4; K 1e7; T 1e4, 6e8; P; from Senegal to
Madagascar and Ethiopia to Zimbabwe, Comoro Is. At Kilimanjaro
in the lowlands scattered.
Pollination: bees (Dodson, 1966); other vectors are possible.
Tabernaemontana stapfiana Britten (Plate 4, 53 & Plate 5, 1e3).
Shape: spheroidal, circular ~40 mm.
Apertures: 3e4 colporate, colpus length max 1/4 to 1/3 and
narrow, pori broad rectangular, mesopori vaulted and granular.
Exine: tectate, scabrate.
Habitus: tree 5e25 (35) m high.
Habitat: forest; 1400e2300 m. At Kilimanjaro mainly in lower
montane riverine, Cassipourea and Ocotea forests, from 1280 to
2330 m.
Distribution: U 2, 3; K 3e7; T 2, 3, 6e8; DR Congo-Kinshasa,
Rwanda, Burundi, Malawi, Mozambique, Zimbabwe. At Kili-
manjaro in the lower forest belt widespread.
Pollination: bees (Moura et al., 2011); other vectors are possible.
5.2.6. Aquifoliaceae
Trees or shrubs, evergreen or deciduous. Aquifoliaceae today
only include the genus Ilex (Judd et al., 2010), which is composed of
almost 600 species. Although cosmopolitan, it is very unevenly
distributed in terms of the number of species occurring in each
continent. Ilex occurs mostly in the tropics but extends into
temperate regions with oceanic climates. There is only one species
in sub-Saharan Africa (Loizeau et al., 2005).
5.2.6.1. Pollination. Insect-pollinated (Loizeau et al., 2005).
5.2.6.2. Pollen morphology. pollen grains are 3-colporidate and
spheroidal-prolate. Pollen surface is conspicuously gemmate/
clavate (Erdtman, 1952).
Ilex mitis (L.) Radlk. var mitis (Plate 5, 4e6).
Shape: subprolate-spheroidal, triangular (convex) ~35 mm.
Apertures: 3 colpor(id)ate, long broad colpi (5/6), pori badly
defined.
Exine: clavate, pilate with pycnic pila which are smaller towards
the apertures.

Plate 5. APOCYNACEAE: 1e3 Tabernaemontana stapfiana, AQUIFOLIACEAE: 4e6 Ilex mitis var. mitis, ARALIACEAE: 7e8 Cussonia arborea, 9e11 Polyscias fulva, 12e15 Schefflera
myriantha, 16e18 Schefflera volkensii, ASPARAGACEAE: 19e20 Asparagus racemosus, ASTERACEAE: 21e22 Artemisia afra, 23e25 Bidens pilosa, 26e28 Bothriocline amplifolia, 29e31
Bothriocline longipes, 32e33 Carduus keniensis, 34e36 Centaurea praecox, 37e39 Cineraria deltoidea, 40 Conyza newii.
328 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
Habitus: evergreen shrub, or more usually a tree, 4e24 (40) m
tall and up to 0.3 (1) m in diameter at the base of the trunk.
Habitat: upland rain-forest, dry evergreen forest and derived
thicket; 900e2850 (3150) m. At Kilimanjaro mainly in lower and
middle montane riverine, Cassipourea, Ocotea forests as well as
upper montane Podocarpus and Juniperus forests, from 1460 to
3000 m.
Distribution: U 1e3; K 3e6; T 2, 3, 7, 8; Sierra Leone, Nigeria,
Cameroon Republic and Fernando Po to Ethiopia, Eritrea, south to
Cape Province of South Africa, also Madagascar. At Kilimanjaro in
the forest belt widespread.
Pollination: insects (Loizeau et al., 2005), mainly flies (Knuth
et al., 2011).
5.2.7. Araliaceae
Trees or ‘arborescent’, or shrubs or lianas, or herbs. Perennial,
self supporting, or epiphytic, or climbing. Also temperate but
mainly sub-tropical to tropical. Widespread, but especially Indo-
malaya and tropical America. About 700 species in 49 genera
(Watson and Dallwitz, 1992 onwards).
5.2.7.1. Pollination. Insect pollinated (Willemstein, 1987).
5.2.7.2. Pollen morphology. Pollen grains 3-colporate (occasionally
2- or 4-colporate, 3-colpate, or 6-colpate), oblate spheroidal to
prolate (longest axis usually 30e40 mm). Sexine in most cases as
thick as nexine, often thicker at poles than at equator, usually
reticulate (Erdtman, 1952).
Cussonia arborea Hochst. ex A. Rich (Plate 5, 7e8).
Shape: subprolate ~20  27 mm, trilobate ~20 mm.
Apertures: 3-colporate, colpi long (3/4) and narrow, pori rect-
angular broad.
Exine: tectate, sexine not or only slightly thicker than nexine,
exine not thicker at poles, reticulate, heterobrochate.
Habitus: a tree up to 13 m tall, with a bole up to 1 m or more in
diameter.
Habitat: woodland, grouped-tree grassland; 300e2470 m. At
Kilimanjaro in dry savanna forests together with Stereospermum
kunthianum, Ozoroa insignis and Pleurostylia africana, from 1100 to
1230 m.
Distribution: U 1, 3, 4; K 2e5, 6; T 1, 2, 4e8; Ethiopia, Sudan, DR
CONGO, Zambia, Zimbabwe. At Kilimanjaro in the savanna rare.
Pollination: insect. No further details known.
Polyscias fulva (Hiern) Harms (Plate 5, 9e11).
Shape: subprolate ~17  24 mm, triangular convex e trilobate
~17 mm.
Apertures: 3-colporate, colpi (4/5) narrow, pori lalongate and
butterfly shaped.
Exine: tectate, reticulate, dense reticulum, homobrochate, exine
not thicker at poles.
Habitus: a tree to 30 m tall, often with a large grey extremely
straight unbranched cylindrical bole of up to 15 m tall and 1 m in
diameter, generally dividing into a small number of main branches
which themselves each branch in a similar manner.
Habitat: upland and lowland rain-forest, riverine forest, also
upland grassland; 1180e2160 m. At Kilimanjaro a pioneer tree
mainly in lower montane Ocotea and riverine forests, from 1320 to
2090 m.
Distribution: U 2e4; K 5, 7; T 1e4, 6, 7; widely spread from
Guine e Republic to Ethiopia, and southwards through the DR Congo
Republic to Malawi, Zambia, Zimbabwe and Angola. At Kilimanjaro
widespread in the lower forest belt, mainly at the southern slope.
Pollination: insects (Lemmens et al., 2009).
Schefflera myriantha (Bak.) Drake (Plate 5, 12e15) SYN Schefflera
congesta De Wild.
Shape: prolate ~21  29 mm, trilobate ~21 mm.
Apertures: 3-colporate, long (5/6) and narrow colpi, pori
lalongate and butterfly shaped.
Exine: tectate, reticulate, heterobrochate, exine thicker at poles.
Habitus: a robust liane or tree of up to 16 m, or a shrub (Cannon,
1978).
Habitat: moist bamboo thickets, upland rain-forest;
1540e2770 m. At Kilimanjaro in moist lower to middle montane
riverine and Ocotea forests, from 1600 to 2800 m.
Distribution: U2; K ?3, 4, ?6, 7; T2-7; DR CONGO, Ethiopia,
Malawi. At Kilimanjaro widespread in the lower and middle forest
belt of the southern slope.
Pollination: insects. No details known.
Schefflera volkensii (Plate 5, 16e18).
Shape: prolate ~19  29 mm, trilobate.
Apertures: 3-colporate, long (5/6) and narrow colpi with thin
margo, pori elliptic broad to butterfly shaped.
Exine: tectate, reticulate, heterobrochate, exine not (much)
thicker at poles.
Habitus: scandent shrub or tall e sometimes spreading and
much branched e tree up to 24e30 m tall, or an epiphyte upon
other trees.
Habitat: upland rain-forest, upland dry evergreen forest;
1600e3200 m. At Kilimanjaro widespread in lower and middle
montane riverine, Cassipourea and Ocotea forests as well as upper
montane Podocarpus and Juniperus forests, from 1600 to 3170 m.
Distribution: U 3, K 3e6; T 2; Ethiopia. At Kilimanjaro in the
forest belt widespread.
Pollination: insects (bees). No details known.
5.2.8. Asparagaceae
Erect or scandent, mostly glabrous herbaceous or subshrubby
perennials with persitant, evergreen or anually withering branches
growing from a compact or creeping rhizome or rarely a tuber. Only
one genus with a controversial number of species (170e300?),
throughout Africa, most of Europe, Asia and Australia (Kubitzki and
Rudall, 1998).
Pollination: although Hymenoptera and Diptera have been
observed visiting the flowers, the pollination process has not yet
been documented (Kubitzki and Rudall, 1998).
Pollen Morphology: grains are monosulcate, tenui-exinous and
have a psilate-microperforate or slightly reticulate exine; sexine as
thick as nexine or thinner (Erdtman, 1952).
Asparagus racemosus Willd (Plate 5, 19e20).
Shape: prolate ~20  32 mm, ± circular.
Apertures: monosulcate, sulcus (3/4 to 4/5) mostly broad.
Exine: tenui-exinous (pattern ± obscure), sexine as thick as
nexine or thinner, scabrate, microreticulate.
Habitus: climbing shrub to 7 m high.
Habitat: forests, forest margins, secondary forest, wooded
grassland, secondary bushland and grassland; (350-)
1550e2900 m. At Kilimanjaro, in the whole forest belt from dry
colline savanna forest up to upper montane Podocarpus and Juni-
perus forest, also in Chagga Homegardens; 1000e2900 m.
Distribution: U 1e4; K 1, 3e6; T 1e4; Sudan, Ethiopia, Eritrea,
Somalia, Zimbabwe, Asia. At Kilimanjaro widespread.
Pollination: probably insects (Kubitzki and Rudall, 1998).
5.2.9. Asteraceae
Herbs, annual, biennial or perennial, subshrubs, shrubs or less
often trees, sometimes lianas, usually terrestrial, rarely epiphytic or
aquatic, sometimes succulent. The largest family of flowering
plants, cosmopolitan except for Antarctica, with over 1600 genera
and 23,000 species (excluding apomictic microspecies), especially
well represented in grassland, wooded grassland and montane
 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
vegetation, comparatively few in humid tropical lowland forests
(Anderberg et al., 2009).
5.2.9.1. Pollination. The majority of Asteraceae are adapted to
specific pollinators. The most important pollinators are solitary
bees; hummingbirds, sunbirds, other long-tongued insects and flies
are also known as pollination agents (Anderberg et al., 2009, and
literature herein). Wind pollination occurs in the Anthemideae (e.g.
Artemisia).
5.2.9.2. Pollen morphology. pollen in Asteraceae is 3-colporate or
sometimes triporate, usually echinate or spinulate, generally
spheroidal with a transverse furrow in the endexine perpendicular
to each colpus and a tectate ectexine. The ectexine generally con-
sists of a foot layer, columellae and a tectum. The great variation in
pollen form and structure in Asteraceae provides much taxonom-
ically valuable information (Stix, 1960; Skvarla et al., 1977;
Gustafsson and Bremer, 1995).
Artemisia afra Willd (Plate 5, 21e22).
Shape: subprolate-spheroidal, circular, ~25 mm.
Apertures: 3-colporate, pori tall elliptic, colpi long (5/6) and
narrow.
Exine: tectate, distinctly double-columellate, micorechinate/
verrucate, wall thickness even/thinner on poles.
Habitus: woody herb, perennial, 0.3e2 (2.4) m tall.
Habitat: montane to alpine grassland and heathland, at lower
altitudes in grassland and bushed or wooded grassland, also in
secondary vegetation; a pioneer after fires, and may be locally
common or thicket-forming; 1500e4050 m. At Kilimanjaro mainly
in subalpine Erica trimera bushland with Stoebe kilimandscharica,
Anthospermum usambarense and Adenocarpus mannii, below the
forest belt very rarely in thickened edges, from 1400 to 3200 m.
Distribution: U 1, 3; K 2e7; T 2e4, 7; S. DR CONGO, Ethiopia,
Angola, Zambia, Malawi, Mozambique, Zimbabwe, Namibia, South
Africa. At Kilimanjaro scattered in the lower subalpine zone.
Pollination: all Artemisia species are wind pollinated (Wright,
2002).
Bidens pilosa L (Plate 5, 23e25).
Shape: circular, spheroidal, ~41 mm.
Apertures: 3-colporate, circular pori, long narrow colpi.
Exine: tectate, columellate, perforate, echinate, spines with very
acute and long tips.
Habitus: annual herb, erect, 0.1e1.5 m high.
Habitat: common weed of cultivation and ruderal sites, often on
alluvial sites, also in forest margins and grassland in swamps and
along streams. May be locally common. 0e60 and 750e2500 m. At
Kilimanjaro synanthrop in ruderal vegetation (Chagga home-
gardens, roadsides, fallow arable fields and waste places) from the
savanna to the lower montane zone, rarely in riverine forests, from
800 to 2050 m.
Distribution: U 1, 2, 4; K 1e6; T 1e8, Z, P. Pan-tropical and sub-
tropical, also extending into some temperate areas. At Kilimanjaro
widespread below the forest belt.
Pollination: insects (Hymenoptera, Diptera, Lepidoptera) (Torres
and Galetto, 2002; Grombone-Guaratini et al., 2004; Pauly, 2015).
Bothriocline amplifolia (O. Hoffm. & Muschl.) M.G. Gilbert (Plate
5, 26e28).
Shape: circular, spheroidal, ~63 mm.
Apertures: 3 (col) porate, apertures not distinct.
Exine: tectum perforatum, columellate, echinolophate, short,
slender spines with acute tip, ridges thick and high, lacunae angular
but irregularly shaped.
Habitus: woody herb or subshrub 0.9e2.4 m tall.
Habitat: montane forest (margins), often with Podocarpus, Ilex
and Nuxia. At Kilimanjaro mainly in middle montane (riverine,
329
Cassipourea, Ocotea) forests and upper montane (Podocarpus,
Hagenia, Juniperus and Erica excelsa) forests, from 1600 to 3170 m.
Distribution: K 3, 4; T 2. Cameroon. At Kilimanjaro in the forest
belt scattered, with low densities.
Pollination: insects.
Bothriocline longipes (Oliv. & Hiern) N.E. Br (Plate 5, 29e31).
Shape: circular, spheroidal, ~43 mm.
Apertures: 3 (col)porate, apertures not well discernable.
Exine: tectum perforatum, columellate, echinolophate, spines
very short, slender and acute, often inlined, lacunae circular to
hexagonal, ridges distinctly lower and thinner than in B. amplifolia.
Habitus: perennial woody herb or shrub (0.3-) 0.7e3 m tall.
Habitat: upland grassland, scattered tree grassland, swamp and
forest margins and clearings, also in disturbed places. May be
common and gregarious. 850e2700 m. At Kilimanjaro mainly in
forest clearings within Cassipourea and Juniperus forests, together
with Urtica massaica and Vernonia lasiopus, from 1890 to 2640 m.
Distribution: U 1e4; K 2e6; T 2, 3, 6. DR Congo (Kinshasa),
Rwanda, Burundi, Sudan, Angola, Zambia, Malawi. At Kilimanjaro
scattered in the forest belt of the northern slope.
Pollination: insects.
Carduus keniensis R.E. Fr. (Plate 5, 32e33).
Shape: circular, spheroidal, ~54 mm.
Apertures: 3-colporate, short (1/2) narrow colpi, lalongate pore.
Exine: tectate, columellate, echinate, spines resemble equilat-
eral triangles, spines' tips rather obtuse.
Habitus: herb to 2.4 m high, scapose or almost acaulescent.
Habitat: moist sites in afro-alpine moorland and heath zone,
gregarious, may form patches. 2950e4570 m. At Kilimanjaro in
subalpine Erica trimera bushland and forest together with Peuce-
danum kerstenii, Euryops dacrydioides and Senecio schweinfurthii,
and in alpine Helichrysum scrub together with Helichrysum newii, H.
citrispinum, Pentaschistis borussica and Senecio telekii, from 3330 to
4330 m.
Distribution: U 2, 3; K 3, 4; T2. Not known elsewhere. At Kili-
manjaro widespread in the subalpine and alpine zone, in low
densities.
Pollination: insects.
Centaurea praecox Oliv. & Hiern (Plate 5, 34e36).
Shape: subprolate-spheroidal, trilobate, ~31 mm.
Apertures: 3-colporate, long (5/6) and broad colpi with acute
ends, pore elliptic broad tapering off lalongate.
Exine: tectate, columellate, exine thinner towards apertures,
microechinate.
Habitus: perennial herb with stems to 50 cm high.
Habitat: grassland or wooded grassland, occasionally in wood-
land. 950e2600 m. At Kilimanjaro in anthropogenic submontane
Hyarrhenia rufa grasslands, together with Gerbera viridifolia, Gnidia
apiculata and Helichrysum nudifolium, from 1270 to 1400 m.
Distribution: U 1; K 2, 3; T 4, 7. West Africa from Mali to Ghana to
NE Africa in Sudan and Ethiopia, DR Congo (Kinshasa), Burundi,
south to Zambia, Malawi, Mozambique, Zimbabwe. At Kilimanjaro
extremely rare.
Pollination: insects.
Cineraria deltoidea Sond. (Plate 5, 37e39).
Shape: spheroidal, circular e triangular, ~28 mm.
Apertures: 3-colporate, long (4/5) and narrow colpi, with
lolongate but very inconspicuous pore.
Exine: tectate, columellate (rather thin), echinate, many rela-
tively short spines broad based with acute tips.
Habitus: perennial herb, erect or more usually scandent or
trailing. 0.15e3 m high or long.
Habitat: grassland, montane forest and forest margins, glades in
bamboo and Hypericum zone, moorland, especially in moist places,
in lava crevices, may be temporarily co-dominant in secondary
330 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386

vegetation; (1100-)2100e4050 m. At Kilimanjaro mainly in upper
montane Hagenia, Juniperus and Erica excelsa forests and subalpine
Erica trimera bushland and forest, from 1710 to 3900 m.
Distribution: U2, 3; K1, 3e6; T2, 4, 6e7; DR Congo (Kinshasa),
Rwanda, Sudan, Ethiopia, Zambia, Malawi, Mozambique,
Zimbabwe, South Africa. At Kilimanjaro widespread in the subal-
pine zone.
Pollination: insects.
Conyza newii Oliv. & Hiern (Plate 5, 40 & Plate 6, 1e2).
Shape: spheroidal, slightly trilobate ~20 mm.
Apertures: 3-colporate, long (5/6) and open colpi with costa,
colpal membrane sub-psilate, pori lalongate but inconspicuous.
Exine: tectate, columellate, echinate, spines with acute tips and
broad base, tectum between spines is perforated to sub-psilate.
Habitus: shrub or woody herb, 0.3e2.5 m high.
Habitat: forest margins and grassy clearing (especially in drier
forest, e.g. Juniperus or Podocarpus type), montane scrub or bush-
land (with Lantana, Erica), especially in rocky sites; (1050-)
1500e3050 m. At Kilimanjaro in lower montane secondary bush-
land and regeneration stages of disturbed forest, in upper montane
Podocarpus, Hagenia, Juniperus and Erica excelsa forests, avoiding
the dense and moist middle montane forests, from 1580 to 3200 m.
Distribution: U 1e3; K 1e7; T 2, 3, 5, 6. DR CONGO, Rwanda,
Ethiopia. At Kilimanjaro rare in the forest belt.
Pollination: insects.
Conyza ruwenzoriensis R.E.Fr (Plate 6, 3e6).
Shape: spheroidal e slightly trilobate, circular, ~25 mm.
Apertures: 3-colporate, long (4/5) and open colpi with costa,
pore circular e elliptic broad (lalongate).
Exine: tectate, columellate, echinate, spines with acute tips and
broad base, more and smaller spines than in C. newii and
C. vernonioides.
Habitus: erect perennial herb, woody herb or shrub, 0.4e2 m
high. Branched distally.
Habitat: upland grassland, afromontane grassland or Erica-
Stoebe bushland; 1800e3800 m. At Kilimanjaro mainly in upper
montane Erica excelsa forest and subalpine Erica bushland and
Festuca obturbans tussock grassland, from 2480 to 3140 m.
Distribution: U 2, 3; K 3e6; T 2, 4, 7; DR CONGO. At Kilimanjaro
scattered in the upper montane and subalpine zone.

Plate 6. ASTERACEAE: 1e2 Conyza newii, 3e6 Conyza ruwenzoriensis, 7e8 Conyza vernonioides, 9e11 Crassocephalum bojeri, 12e13 Crepis oliveriana, 14e16 Dicoma tomentosa, 17e18
Dicrocephala integrifolia, 19e20 Erlangea calycina, 21e23 Gerbera viridifolia, 24e25 Haplocarpha rueppellii, 26e28 Helichrysum argyranthum, 29e30 Helichrysum citrispinum, 31e33
Helichrysum forskahlii var. forskahlii, 34e36 Helichrysum schimperi, 37e38 Hirpicium diffusum.
 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
Pollination: insects.
Conyza vernonioides (A. Rich.) Wild (Plate 6, 7e8).
Shape: spheroidal, circular, ~25 mm.
Apertures: 3-colporate, long (4/5) and open colpi with costa,
pore circular e elliptic broad (lalongate).
Exine: tectate, columellate, echinate, spines with long (some-
times bent) acute tips and broad base, tip seems slightly set off the
base.
Habitus: shrub or small tree, 1.5e9 m high. Much-branched.
Habitat: bamboo zone, Hagenia-Myrsine (Rapanea) zone, Erica-
Stoebe zone, in glades and forest margins, in damp sites, on ridge
crests or steep slopes; 2300e3600 (3750) m. At Kilimanjaro
mainly in upper montane Hagenia, Juniperus and Erica excelsa forest
and subalpine Erica bushland from 2310 to 3920 m.
Distribution: U 2e3; K 3e6; T 2; DR CONGO, Ethiopia. At Kili-
manjaro scattered in the upper montane and subalpine zone.
Pollination: insects.
Crassocephalum bojeri (DC.) Robyns (Plate 6, 9e11) SYN C. bojeri,
Senecio bojeri, Solanecio angulatus.
Shape: subprolate, trilobate, ~37  40 mm.
Apertures: 3-colporate, long and open colpi, colpi straight with
obtuse ends, pore circular to elliptic tall (lolongate).
Exine: tectate, columellate, echinate, many spines short with
acute tips.
Habitus: scrambling perennial succulent herb, occasionally
prostrate, 0.6e3 (10) m long or high.
Habitat: forest, especially in riverine or lakeshore forest, bush-
land, bushed grassland, thickets. (250-) 600e1800 (2500) m. At
Kilimanjaro in savanna riverine forests, submontane Croton-Calo-
dendrum forest, as well as in Chagga homegardens, from 960 to
1600 m.
Distribution: U 1e4; K 1e7; T 1e8. Cameroon, Equatorial
Guinea, Gabon, DR Congo (Kinshasa), Rwanda, Burundi, Sudan,
Ethiopia, Somalia, Angola, Zambia, Malawi, Mozambique,
Zimbabwe, South Africa, Comoro Is., Madagascar, Yemen. At Kili-
manjaro scattered.
Pollination: insects.
Crepis oliveriana (Kuntze) C. Jeffrey (Plate 6, 12e13) SYN Crepis
newii subsp. oliveriana.
Shape: spheroidal to hexagonal with convexly rounded sides,
~42 mm.
Apertures: 3 (col)porate, pori inconspicuous, colpi divided into
three lacunae connected by narrow interlacunar gaps.
Exine: tectate, thin columella, echinolophate, abporal lacunae
large, angular and broader towards the poles, one line of relatively
short spines on each ridge.
Habitus: perennial herb 0.1e1.2 m high.
Habitat: montane grassland, giant heath zone, often on lava
soils. 1650e3750 m. At Kilimanjaro in upper montane Erica excelsa
forests and on rocks in Hagenia riverine forests, from 2700 to
3510 m.
Distribution: K 3, 4; T 1, 2, 7. Cameroon. At Kilimanjaro scattered
in the upper montane and subalpine zone.
Pollination: insects.
Dicoma tomentosa Cass (Plate 6, 14e16).
Shape: subprolate, triangular, ~28  33 mm.
Apertures: 3-colporate, colpi long and narrow (wider at equa-
tor) with costae, pori lalongate/elliptic broad.
Exine: tectate, aerolate, exine very thick, thinner at the poles,
supporting columellae thick, densely distributed and more or less
straight, densely verrucate.
Habitus: erect often much-branched and bushy, rather woody
annual herb 0.5e0.75 m tall.
Habitat: semi-desert grassland and bushland, grassland, decid-
uous bushland and woodland, often on rock outcrops. 400e1550 m.
331
At Kilimanjaro on fallow arable land together with Vernonia kar-
aguensis, Indigofera schimperi var. baukeana and Hibiscus pandur-
iformis at 900 m.
Distribution: U 1; K 1e4, 6, 7; T 1e3, 5e7. Senegal to Sudan,
Ethiopia, Somalia, south to Namibia, South Africa. Egypt, east to
Pakistan and W India. At Kilimanjaro very rare.
Pollination: insects (bees) (Sivaram, 2001).
Dicrocephala integrifolia (L. f.) Kuntze (Plate 6, 17e18).
Shape: spheroidal, trilobate, ~20 mm.
Apertures: 3-colporate, colpi long (5/6), wide at the equator and
tapering towards poles, pore (lolongate?) inconspicuous.
Exine: continuous tectum, columellate, echinate, spines with
long and acute tip which is often slightly bent.
Habitus: annual herb, 0.1e1 m high, erect or procumbent.
Habitat: pioneer in fallow land, a weed of cultivation, in dry or
moist forest margins, in upland grassland, bamboo, bushed or
wooded grassland, often in moist situations, in swamps or on
stream banks; 1100e3000 m. At Kilimanjaro in riverine forests and
Chagga homegardens, from 1210 to 2600 m.
Distribution: U 1e4; K 1e6; T 2, 3, 5e8. Widespread in tropical
and South Africa, Madagascar, the Middle East and tropical Asia. At
Kilimanjaro scattered.
Pollination: insects.
Erlangea calycina S. Moore (Plate 6, 19e20) SYN Bothriocline
calycina (S. Moore) M. G. Gilbert comb. nov.
Shape: spheroidal to subprolate ~50 mm, circular e trilobate.
Apertures: 3-colporate, colpi slightly recessed, pori
inconspicuous.
Exine: tectate, columellate, subechinolophate, spines long and
slender with acute tips, often bent, lacunae of variable shape and
size.
Habitus: perennial herb 0.2e0.75 m tall, rhizotamous. Stems
several, ascending or decumbent.
Habitat: grassland and Acacia grassland, often on black cotton
soils. May form extensive stands. 1350e2400 m. At Kilimanjaro in
savanna grassland together with Athroisma hastifolium, Solanecio
goetzei and Themeda triandra at 1390 m.
Distribution: K 3, 4, 6; T 2. Not known elsewhere. At Kilimanjaro
very rare.
Pollination: insects.
Gerbera viridifolia (DC.) Sch. Bip. (Plate 6, 21e23).
Shape: subprolate, trilobate, ~36  45 mm.
Apertures: 3-colporate, long and narrow colpi, small pori
lalongate.
Exine: tectate, columellate, very thick exine, psilate to
microgranulate.
Habitus: herb with woody rootstock and fleshy roots.
Habitat: a pyrophyte of deciduous woodland, open bushland,
wooded grassland and grassland. 850e2750 m. At Kilimanjaro in
anthropogenic submontane Hyarrhenia rufa grasslands, together
with Centaurea praecox, Gnidia apiculata and Helichrysum nudifo-
lium, from 1270 to 1400 m from 1270 to 1400 m.
Distribution: U 1e4; K 1e6; T 1e8. Cameroon, Sudan, Ethiopia,
Somalia, DR Congo (Kinshasa), Burundi, south tropical and South
Africa. At Kilimanjaro very rare.
Pollination: insects (Teeri et al., 2006).
Haplocarpha rueppellii (Sch. Bip.) K. Lewin (Plate 6, 24e25).
Shape: spheroidal, circular - trilobate.
Apertures: 3-colporate, colpi and pori inconspicuous.
Exine: tectate, two distinct columellate layers, partially caveate
with thread-like columellae spanning it at intervals, echinate,
50e57 spines distributed evenly across the grain. The spines are
2.5e3 mm high and 3e4 mm in diameter at the base, exine between
spines smooth.
Habitus: perennial scapose or occasionally escapose herb.
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Occurring in dense mats with many thick, almost tuberous roots.
Habitat: montane grassland or moorland, at higher altitudes in
sheltered moist sites, also in upper bamboo region. 2550e4650 m.
At Kilimanjaro in subalpine Erica trimera bushland and forest, Carex
monostachya bogs, from 3220 to 4100 m.
Distribution: U 3; K 2e4, 6; T 2. Ethiopia, South Africa. At Kili-
manjaro scattered in the subalpine zone.
Pollination: insects.
Helichrysum argyranthum O. Hoffm (Plate 6, 26e28).
Shape: subprolate ~21  26 mm, trilobate ~21 mm.
Apertures: 3-colporate, recessed colpi long and open, pori
elliptic broad (lalongate) but mostly inconspicuous.
Exine: tectate, columellate, echinate, short, evely distributed
spines with broad base and short acute tip, exine inbetween spines
psilate/scabrate.
Habitus: shrub or shrubby herb, rarely described as scandent,
0.1e1.8 (2.4) m high. With spreading branches.
Habitat: in clearings in bamboo zone, in forest margins, in
grassland and among giant heath in the lower moorlands, towards
the upper part of its range in gorges and gullies; may be common in
bamboo and giant heath zone, especially in Uganda; 2100e3700 m.
At Kilimanjaro in upper montane Erica excelsa and Hagenia forests,
in subalpine Erica trimera bushland, from 2700 to 3700 m.
Distribution: U1-3; K1, 3e6; T2. DR Congo (Kinshasa), Rwanda,
Sudan, Ethiopia. At Kilimanjaro scattered in the upper montane and
subalpine zone.
Pollination: insects.
Helichrysum citrispinum Delile (Plate 6, 29e30).
Shape: spheroidal, circular ~25 mm.
Apertures: 3-colporate, long and wide colpi, pori ± circular but
mostly inconspicuous.
Exine: tectate, columellate, echinate, spines short with broad
base and acute tip, densly distributed.
Habitus: shrub or shrubby herb 0.15e1.2 m high.
Habitat: rocky or stony ground in the afro-alpine zone, in the
lower part of this zone in tussock grassland or among giant heath;
3200e5100 m. At Kilimanjaro in subalpine Erica trimera bushland
and forest, in alpine Helichrysum scrub, from 3120 to 4650 m.
Distribution: U3; K3-5; T2. Ethiopia. At Kilimanjaro widespread
and partly dominant in the subalpine and alpine zone.
Pollination: insects.
Helichrysum forskahlii (J.F. Gmel.) Hilliard & B.L. Burtt var. for-
skahlii (Plate 6, 31e33).
Shape: spheroidal, trilobate.
Apertures: 3-colporate, recessed colpi long (4/5), wide at the
equator with acute ends, pori lalongate with pointy ends.
Exine: tectate, columellate, echinate, spines short with very
broad base and acute tip, exine between spines distinctly granular.
Habitus: erect or sometimes scandent herb or shrub, 0.3e1 m
high.
Habitat: upland and alpine grassland, also in bushland, bamboo,
heath and forest margins, often in boggy sites, but also on dry stony
sites, in secondary situations; may be local common; 1050e4000
(4700) m. At Kilimanjaro below the forest belt in lower montane
anthropogenic Bulbostylis meadows (var. forskahlii), in upper
montane Erica excelsa forest, and subalpine Festuca obturbans tus-
sock grassland and Erica bushland and forest, in alpine Helichrysum
scrub 2730e4650 m.
Distribution: U 1e4; K 1e6; T 1e4, 6e8; from Nigeria to Sudan
and Ethiopia on the north, throughout central Africa to Angola and
Zimbabwe in the south; also in Yemen. At Kilimanjaro widespread
in the subalpine and alpine zone.
Pollination: insects.
Helichrysum schimperi (Sch. Bip. ex A. Rich.) Moeser (Plate 6,
34e36).
Shape: subprolate to spheroidal, trilobate.
Apertures: 3-colporate, colpi long, wide at the equator and
tapering towards poles, pore elliptic tall (lolongate).
Exine: tectate, columellate, rather sparsely distributed spines
with broad base and acute tip, exine inbetween spines distinctly
granular.
Habitus: perennial herb, woody herb or shrublet. Erect with
scrambling branches to 3.6 m high, or the whole plant decumbent
and straggling.
Habitat: montane grassland, secondary bushland, riverine for-
est, dry forest margins, giant heath zone, often in rocky sites; (900-)
1350e3300 m. At Kilimanjaro at forest edges and clearings, in
lower montane disturbed forest, in upper montane Hagenia and
Erica excelsa forest, from 1620 to 3310 m.
Distribution: U 1e4; K 1, 3e6; T 2, 3, 6e8. DR Congo (Kinshasa),
Rwanda, Sudan, Ethiopia, Socotra, Zambia, Malawi, Mozambique,
Zimbabwe. At Kilimanjaro in the forest belt scattered.
Pollination: insects.
Hirpicium diffusum (O. Hoffm.) Roessler (Plate 6, 37e38).
Shape: spheroidal, circular.
Apertures: 3-colporate, colpi broad with acute ends but incon-
spicuous, pori circular to elliptic broad (lalongate).
Exine: tectate, psilate, lophate, single columellate layer with the
columellae aggregated under lophae, the lophae sometimes over-
lap one another.
Habitus: annual or short-lived perennial herb, (5-) 30e80 cm
tall. Much-branched, often prostrate or sometimes shrubby with
long woody tap-root.
Habitat: grassland, black cotton soils, plains with poor soils and
scattered herbs, wooded grassland, woodland, stony eroded hill-
sides, rock outcrops, river bank, disturbed sites. 500e3000 m. At
Kilimanjaro in savanna grasslands and anthropogenic submontane
Hyarrhenia rufa grasslands, from 1000 to 1800 m.
Distribution: U 1; K 1e7; T 1e5, 7. Ethiopia. At Kilimanjaro in the
lower parts of the mountain below the forest belt scattered.
Pollination: insects.
Lactuca capensis Thunb (Plate 7, 1e2). SYN L. inermis.
Shape: spheroidal, circular, hexagonal ~40 mm.
Apertures: 3-colporate, colpi divided into three lacunae, con-
nected by narrow interlacunar gaps, pori inconspicuous.
Exine: tectate, echinolophate, abporal lacunae are angular;
paraporal lacunae pentagonal, spines short with broad base and
acute tip, polar regions each with a single row of spines bordering
the adjacent lacunae.
Habitus: perennial herb 0.05e1.5 (2.4) m high. Erect with a
long thick taproot.
Habitat: grassland and a weed of cultivation, on fallow land, in
roadsides. (650-) 1000e2800 (3450) m. At Kilimanjaro in grass-
lands and meadows from the savanna to the lower forest border in
the lower montane zone, from 920 to 1880 m.
Distribution: U 1e4; K 1e6; T 1e8. At Kilimanjaro below the
forest belt widespread.
Pollination: insects.
Pluchea nitens O. Hoffm (Plate 7, 3e5). SYN Nicolasia nitens.
Shape: spheroidal, circular.
Apertures: 3-colporate, colpi wide with costae and of medium
length (3/5), pori ± circular to slightly lalongate inconspicuous.
Exine: tectate, columellate, echinate, densely distributed long
spines with borad base and acute tips, extending columellae into
the spines well visible.
Habitus: annual or short-lived perennial herb 0.05e0.75 m high.
Habitat: within the dry bushland zone in seasonal rivers and
lakes or in beds of temporary pools on lake shore flats, but also on
alkaline soils on volcanoes; on sandy, volcanic, saline or alkaline
soils; may be locally common and forms dense mats among
 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386 333

Plate 7. Asteraceae: 1e2 Lactuca capensis, 3e5 Pluchea nitens, 6e8 Schkuhria pinnata, 9e12 Senecio cyaneus, 13e16 Senecio johnstonii, 17e18 Siegesbeckia abyssinica, 19e20
Sphaeranthus bullatus, 21e23 Spilanthes mauritiana, 24e26 Stoebe kilimandscharica, 27e29 Tagetes minuta, 30e32 Tolpis capensis.

grasses; (300-) 600e1800 m. At Kilimanjaro in grassland on alka- Pollination: insects.

line soil at 950 m.
Distribution: K 1, 3, 4, 6, 7; T 1, 2, 5, 7. Zambia, Zimbabwe,
Botswana, Namibia. At Kilimanjaro extremely rare.
Pollination: insects.
Schkuhria pinnata (Lam.) Kuntze ex Thell (Plate 7, 6e8).
Shape: spheroidal, circular e slightly trilobate ~28 mm.
Apertures: 3-colporate, colpi long and narrow, pori lalongate
with pointy ends.
Exine: tectate, columellate, echinate, spines with broad, merged
bases and acute tips, columellae extending into broad part of
spines.
Habitus: Annual herb 0.05e0.5 m high. Much-branched.
Habitat: Ruderal sites, gardens, arable land, occasionally in dry
woodland, grassland or bushland. May form almost pure stands.
950e1950 m. At Kilimanjaro a weed on fallow arable land, at 930 m.
Distribution: U 1; K 3e6; T2. Ethiopia, Zimbabwe, Mozambique,
Botswana, originally from America but now a widespread. At Kili-
manjaro rare.
Senecio cyaneus O. Hoffm (Plate 7, 9e12).
Shape: subprolate, trilobate.
Apertures: 3-(zono)colporate, recessed colpi long and wide,
tapered at ends, pori elliptic tall (lolongate).
Exine: tectatum perforate, columellae with thin, short and un-
branched rods, sexine thicker than nexine, echinate, densely
distributed spines short with broad base and acute tip.
Habitus: perennial ± erect weak-stemmed herb or straggling
woody herb, 0.3e1.8 m high.
Habitat: upland grassland and heath zone, less often in forest
margin; 2300e3350 m. At Kilimanjaro mainly in upper montane
Hagenia (riverine) and Erica excelsa forest from 2500 to 3500 m.
Distribution: T2, 6. Endemic to Kilimanjaro, Mt. Meru, Hanang
and the Lukwangule Plateau. At Kilimanjaro in the upper part of the
forest belt scattered.
Pollination: species of the genus Senecio are entemophilous
(flies and other insects) (Niemel€ a and Pellikka, 2004).
Senecio johnstonii Oliv (Plate 7, 13e16). SYN Dendrosenecio
334 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
kilimanjari (Mildbr.) E.B. Knox.
Shape: subprolate - prolate, trilobate.
Apertures: 3-colporate, recessed colpi long and open, tapered at
ends, large pori lolongate.
Exine: tectatum perforate, columellate, sexine thicker than
nexine, echinate, sparsely distributed, very short (micro) spines
slim with acute tip.
Habitus: upright, polycarpic plant to 10 m tall.
Habitat: moist sites in the afro-alpine zone, particularly along
streams, larger ravines and bogs; 2940e4280 m.
Distribution: T 2. Endemic to Kilimanjaro.
Pollination: species of the genus Senecio are entemophilous
(flies and other insects) (Niemela € and Pellikka, 2004).
Siegesbeckia abyssinica (Sch. Bip. ex Walp.) Oliv. & Hiern (Plate 7,
17e18) SYN Micractis bojeri.
Shape: subprolate e spheroidal, circular.
Apertures: 3 (zono) colporate, colpi long and open with tapered
ends, circular pori inconspicous.
Exine: tectate, echinate, columellae extending into the base of
spines, long and slim spines with broad base and acute tip which is
regularly bent.
Habitus: annual or sometimes short-lived perennial herb
0.3e2.5 m high.
Habitat: streamsides in grassland or forest, seasonally flooded
grassland, swamp grassland. 1350e2750 m. At Kilimanjaro in lower
montane forest clearings together with Ipomoea involucrata, Tha-
lictrum rhynchocarpum and Acalypha psilostachya, from 2100 to
2200 m.
Distribution: U 2, 3; K 3e6; T 1e4, 7, 8. From Nigeria to Ethiopia
and south to DR Congo and Malawi, Madagascar. At Kilimanjaro
very rare.
Pollination: insects.
Sphaeranthus bullatus Mattf (Plate 7, 19e20).
Shape: spheroidal, subprolate ~19  23 mm.
Apertures: 3-colporate, colpi long and narrow, slightly recessed,
pori lalongate.
Exine: tectate, columellate, echinate, loosly distributed spines
short and resembling equilateral triangles in shape.
Habitus: annual or perennial herb, 0.2e1 m tall. Much-
branched, erect or decumbent and rooting where touching the
ground.
Habitat: moist sites along streams, lake edges, rock potholes,
swampy sites, in seasonally moist grassy plains or along seasonal
pools, often on impeded drainage soils, may be locally common or
form mats; 50e1900 m. At Kilimanjaro in swamps, streamsides and
lake shores of the savanna and submontane zone, together with
Spilanthes mauritiana, Leersia hexandra and Commelina diffusa, as
well as ruderal vegetation on roadsides, from 930 to 1600 m.
Distribution: K 3, 4, 6; T 2, 3, 5; not known elsewhere. At Kili-
manjaro below the forest belt scattered.
Pollination: insects.
Spilanthes mauritiana (A. Rich. ex Pers.) DC (Plate 7, 21e23). SYN
Acmella caulirhiza.
Shape: circular, (sub) prolate.
Apertures: 3 col(por)ate, colpi long, slightly open and sunken in,
pori inconspicuous.
Exine: tectate, columellate, exine thick, echinate, short blunt
spines with broad base.
Habitus: perennial or annual herb, decumbent or scrambling, up
to 0.15 m high or 0.6 m long.
Habitat: swampy or seasonally wet sites, river-banks, cultivated
areas, forest margins. 750e3100 m. At Kilimanjaro in swamps,
streamsides and lake shores of the savanna and submontane zone,
together with Sphaeranthus bullatus, Leersia hexandra and Com-
melina diffusa, as well as in wet grasslands and moist ruderal
vegetation (Chagga homegardens, roadsides, trampled ground)
from the savanna to the lower montane zone, from 790 to 2050 m.
Distribution: U 1e4; K 1e7; T 1e8; widespread in tropical and
subtropical Africa and Madagascar. At Kilimanjaro below the forest
belt widespread.
Pollination: insects, e.g. by carpenter bees Xylocopa (Pauly,
2015).
Stoebe kilimandscharica O. Hoffm (Plate 7, 24e26).
Shape: circular to trilobate, spheroidal ~23 mm.
Apertures: 3-colporate, colpi long, slightly open and sunken in,
pori ± circular.
Exine: tectate, columellate, echinate, short blunt spines with
broad base.
Habitus: shrub, 1.2e6 m high. Much-branched.
Habitat: heath zone, moorland, may be locally (co-) dominant,
either on volcanic soil or on moist ground, also in bamboo and
lower forest zone; (1950-) 2400e3900 m. At Kilimanjaro mainly in
subalpine Erica trimera bushland with Artemisia afra, Antho-
spermum usambarense and Adenocarpus mannii, from 2530 to
3660 m.
Distribution: U1, 3; K2-4, 6; T2, 7; DR Congo (Kinshasa), Malawi.
At Kilimanjaro in the subalpine zone scattered.
Pollination: insects.
Tagetes minuta L (Plate 7, 27e29).
Shape: circular to seemingy trilobate (due to thick tectum),
circular, ~45 mm.
Apertures: 3-colporate, colpi long and wide with acute ends,
pori lalongate/rectangular broad.
Exine: tectate, columellate, tectum very thick and perforate,
echinate, spines base broad but not merged, tips acute and long.
Habitus: annual herb, 0.1e2.5 m high.
Habitat: weed of cultivation and post-cultivation, in ruderal
sites. May occur in dense patches from (850-) 1300e2750 m. At
Kilimanjaro in ruderal vegetation (Chagga homegardens, roadsides,
fallow arable fields and waste places) from the savanna to the lower
montane zone, here also in wet grasslands, from 820 to 1930 m.
Distribution: U 1e3; K 1e7; T 1e3, 5, 7, 8; originally from South
America but now a widespread weed in Africa, S. Europe, S. Asia and
Australia. At Kilimanjaro below the forest belt widespread.
Pollination: insects (Basilio et al., 2006).
Tolpis capensis (L.) Sch. Bip (Plate 7, 30e32).
Shape: spheroidal, circular, ~32 mm.
Apertures: 3 (col)porate, apertures not clear, short colpi and/or
elliptic tall pori.
Exine: tectate, tectum very thick, columellate, echinolophate,
lacunae regularly spaced, spines base broad and merged, tips acute
and long.
Habitus: perennial scapose herb. Taproot fleshy and stout.
Habitat: grassland, especially on thin or rocky soils and where
regularly burned, also in open woodland and as a weed of culti-
vation. May be locally common. 1800e3300 m. At Kilimanjaro
mainly in wet anthropogenic vegetation (meadows, trampled
ground) in the lower montane zone, from 1450 to 2130 m.
Distribution: U 1; K 3, 4; T 2e4, 7. DR Congo (Kinshasa), Rwanda,
Burundi, Sudan, Angola, Zambia, Malawi, Mozambique, Zimbabwe,
South Africa, Madagascar. At Kilimanjaro below the forest belt
scattered.
Pollination: insects.
Tridax procumbens L (Plate 8, 1e3).
Shape: spheroidal, circular ~36 mm.
Apertures: 4-colporate, very short oval colpi, lalongate/elliptic
broad pori.
Exine: tectate, columellate, echinate, spines with broad base and
acute often bent tip.
Habitus: annual or perennial herb, sprawling or ascending but
 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386 335

Plate 8. ASTERACEAE: 1e3 Tridax procumbens, 4e6 Vernonia brachycalyx, 7e8 Vernonia colorata, 9e10 Vernonia lasiopus, 11e12 Vernonia syringifolia, BALANITACEAE (Zygophylla-
ceae): 13e15 Balanites aegyptiaca, 16e17 Balanites maughamii spp. acuta, BALSAMINACEAE: 18e19 Impatiens pseudoviola, 20e21 Impatiens walleriana, Basellaceae: 22e24 Basella
alba, BEGONIACEAE: 25e26 Begonia johnstonii, BIGNONIACEAE: 27e29 Jacaranda mimosifolia, 30e31 Markhamia lutea.

with erect inflorescences, 0.15e0.5 m tall.
Habitat: weed of cultivation, also on waste ground, an invader of
bare soil. May form patches or mats. 0e1700 (2850) m. At Kili-
manjaro on rocks (together with Actiniopteris dimorpha), in grass-
lands and in ruderal vegetation (Chagga homegardens, roadsides,
fallow arable fields and waste places) in the lower areas (colline-
submontane), from 800 to 1440 m.
Distribution: U 1e4; K 1, 3e7; T 1e8; Z. Originally from Central
America, now a pantropical weed. At Kilimanjaro widespread.
Pollination: insects (Varalakshmi and Raju, 2013).
Vernonia brachycalyx O. Hoffm (Plate 8, 4e6).
Shape: circular to trilobate, spheroidal ~47 mm.
Apertures: 3-colporate, narrow and broad polar areas.
Exine: tectate, micropunctate tectum, collumelate, sub-
echinolophate, spines long and pointy on the ridges, not merged at
base.
Habitus: erect or ± scrambling or scandent weakly woody shrub
or rarely small tree 1.5e6 m tall.
Habitat: dry forest margins, riverine forest and thicket margins,
also in secondary bushland derived from forest. May form pure
stands. 750e2700 m. At Kilimanjaro mainly in forest clearings and
forest edges within Croton-Calodendrum and Cassipourea forests,
from 1430 to 1820 m.
Distribution: U 1e4; K 1, 3e7; T 1e7. DR Congo (Kinshasa),
Rwanda, Ethiopia, Angola, Zambia, Malawi. At Kilimanjaro rare.
Pollination: Hymenoptera, Lepidoptera (Jones, 1966).
Vernonia colorata (Willd.) Drake (Plate 8, 7e8).
Shape: spheroidal, circular ~49 mm.
Apertures: 3-colporate, colpi and pori indistinct.
Exine: tectate, collumelate, echinolophate, lacunae roundish
and regularly spaced, spines slim and relatively short with blunt
tips, spines base not merged.
Habitus: shrub or small tree (0.75-) 1.5e7.5 m tall.
Habitat: Brachystegia-Isoberlinia woodland, riverine or lakeside
vegetation. 1e1200 m. At Kilimanjaro in riverine and groundwater
forests of the colline zone, also a weed in homegardens, from 780 to
1080 m.
Distribution: K 7; T 1, 2, 3, 4, 6, 8. W and S tropical Africa, DR
Congo (Kinshasa). At Kilimanjaro rare.
Pollination: insects.
Vernonia lasiopus O. Hoffm (Plate 8, 9e10).
Shape: circular to trilobate, spheroidal ~49 mm.
336 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
Apertures: 3-colporate, position of colpi visible due to high
ridges, pori indistinct.
Exine: tectate, columellate, micropunctate, echinolophate,
ridges high, lacunae roundish and regularly spaced, spines slim and
relatively short with acute tips, spines base not merged.
Habitus: woody herb or shrub 0.4e3 m tall, rarely climbing to
4.5 m.
Habitat: forest clearings, forest margins, secondary bush derived
from forest, riverine thicket, secondary grassland in forest or dry
bush zone, roadsides. May be abundant in abandoned cultivation.
1050e2650 m. At Kilimanjaro mainly in forest clearings within
submontane and lower montane Croton-Calodendrum and Cassi-
pourea forest, together with Urtica massaica and Bothriocline long-
ipes, also a weed in Chagga homegardens and coffee plantations,
from 1070 to 2000 m.
Distribution: U 2e4; K 1, 3, 4, 6, 7; T 1e3. Rwanda, Sudan,
Ethiopia. At Kilimanjaro scattered.
Pollination: Hymenoptera, Lepidoptera (Jones, 1966).
Vernonia syringifolia O. Hoffm (Plate 8, 11e12).
Shape: spheroidal, circular ~47 mm.
Apertures: 3-colporate, colpi narrow, pori lolongate/elliptic tall.
Exine: tectate, micropunctate, columellate, subechinolophate,
spines short with broad base and acute tip.
Habitus: often ± scrambling or scandent woody herb or weak
shrub 0.8e4.5 m tall.
Habitat: moist forest and forest margins, secondary bush or
woodland in the forest zone. May be locally dominant in secondary
situations. 1550e3050 m. At Kilimanjaro secondary bushland,
clearing in submontane and lower montane forest, together with
Cyathula cylindrical, Plectranthus albus and Thalictrum rhyncho-
carpum, from 1480 to 2380 m.
Distribution: U 2, 3; K 1, 3e6; T 2, 4, 7. DR Congo (Kinshasa),
Rwanda, Sudan, Zambia, Malawi. At Kilimanjaro rare.
Pollination: Hymenoptera, Lepidoptera (Jones, 1966).
5.2.10. Balanitaceae (today part of Zygophyllaceae).
Trees, shrubs, subshrubs or annual or perennial herbs, often
with joined branches and swollen at the nodes. Comprising 22
genera and 230e240 species in hot dry regions of Europe, Asia,
Australia, Africa and the Americas (Sheahan, 2007).
5.2.10.1. Pollination. Flowers are generally insect-pollinated
(Sheahan, 2007).
5.2.10.2. Pollen morphology. Pollen grains are eurypalynous and
heterogeneous, even at subfamily level. In most of the family, pol-
len grains are 3-colpate, 3-colporate or tricolporidate, rarely 6-
rugorate. Exine thickness varies in the range 1e8 mm, the sexine
is usually of the same thickness as the nexine but may be thicker,
e.g. in Seetzenia. Exine ornamentation is mainly reticulate, occa-
sionally also described as rugulate, baculate and retipilate. The
grains vary in shape from prolate to oblate-spheroidal, they vary
also in size, the longest axis from 10 to 67 mm (Erdtman, 1952;
Sheahan, 2007).
Balanites aegyptiaca (L.) Delile (Plate 8, 13e15)
(Zygophyllaceae).
Shape: spheroidal-trilobate, subprolate ~31  39 mm.
Apertures: 3-colporate, large pori elliptic tall to circular with
annulus, colpi long (4/5) and narrow.
Exine: tectate, sexine thicker than nexine, equatorial exine thin,
thickened around apertures, columellate, perforate, (striato)-
reticulate.
Habitus: Semi-evergreen or sometimes deciduous shrub or
small tree up to 12 (15) m high, extremely variable in many of its
characters. Bole usually straight, often fluted, branches spreading
irregularly or pendulous, sometimes forming a rounded crown.
Habitat: Deciduous bushland, scattered tree grassland, wooded
grassland, thicket; 800e1700 m. At Kilimanjaro in savanna wood-
land from 1010 to 1390 m.
Distribution: K 3, 4, 6; T 1e3; throughout much of Africa from
Mauritania to Somalia and from Egypt southwards to Zambia and
Zimbabwe, W. Arabia and the Jordan valley. At Kilimanjaro in the
savanna scattered.
Pollination: insects: Hymenoptera, Diptera, Coleoptera (Ndoye
et al., 2004).
Balanites maughamii Sprague ssp. acuta Sands (Plate 8, 16e17)
SYN: B. wilsoniana (Zygophyllaceae).
Shape: spheroidal ~36 mm, circular to trilobate.
Apertures: 3-colporate, large pori circular (Ø 8e9 mm) to elliptic
tall with annulus, colpi long (4/5), broader at equator and tapering
towards poles.
Exine: tectate, columellate, perforate, finley reticulate,
homobrochate.
Habitus: Deciduous or semi-deciduous tree up to 20 (25) m
high, with a spreading, rounded crown, rarely a low shrub 1.5e2 m
high; trunk fluted.
Habitat: Evergreen coastal forest, coastal or riverine thicket,
groundwater forest; 0e1000 m. At Kilimanjaro in savanna riverine
forest at 1000 m.
Distribution: K 4, 7; T 2, 3, 6, 8; not known elsewhere. At Kili-
manjaro very rare.
Pollination: insects; no further details known.
5.2.11. Balsaminaceae
Annual or perennial herbs, sometimes with tubers or rhizomes,
occasionally subshrubs. Two genera (one of which monotypic) and
ca. 1000 species in Europe, Africa, Asia, North and Central America
(Fischer, 2004).
Pollination: pollination biology and breeding system of the
majority of species have not yet been studied, and most available
studies have been conducted with the temperate species Impatiens
capensis and I. pallida. Published reports and personal field obser-
vations suggest the following animals as possible pollinators:
bumblebees, solitary bees, and butterflies (e.g. I. walleriana, I.
hoehneliana) (Fischer, 2004 and literature herein).
Pollen morphology: pollen grains are 4-colpate and rectangular
in the North American and Eurasion taxa, 4-colpate rectangular, 4-
colpate square or 3-colpate triangular in the taxa from tropical
Africa and Asia (Huynh, 1968; Lu, 1991). Pollen grains are flattened,
reticulate and the sexine is slightly thicker than nexine (Erdtman,
1952).
Impatiens pseudoviola Gilg (Plate 8, 18e19).
Shape: triangular convex to spheroidal ~36 mm, circular.
Apertures: 3-colpate, angulaperturate, colpi very short.
Exine: reticulate, heterobrochate, simplicolumellate, brochi
without columellae.
Habitus: annual or short-lived perennial to 30e40 cm tall;
stems erect to decumbent, simple or moderately branched.
Habitat: damp shaded upland rain-forest, often on mossy banks,
fallen branches or tree stumps and by waterfalls; 1550e3200 m. At
Kilimanjaro mainly in lower to middle montane moist forests
(riverine, Cassipourea and Ocotea forest) from 1240 to 2920 m.
Distribution: K3, 4, 7; T 2, 3, 5, 6; not known elsewhere. At
Kilimanjaro in the forest belt widespread.
Pollination: various insects and wind (Mansor et al., 2004).
Impatiens walleriana Hook f (Plate 8, 20e21).
Shape: rectangular ~29  47 mm, oblate.
Apertures: 4-colpate, very short colpi (max 1/6).
Exine: reticulate, heterobrochate, lumen larger at poles, sim-
plicolumellate, brochi without columellae.
 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
Habitus: rather succulent perennial 0.3e0.7 (0.8) m tall.
Habitat: in damp, often shaded, places in upland and coastal
rain-forest, particularly in riverine thickets, gullies and damp rocky
places; 0e2000 m. At Kilimanjaro in colline-submontane riverine
forests, humid shaded embankments of irrigation funnels and road
ditches in the Chagga homegardens, inside Chagga homegardens,
from 1080 to 1770 m.
Distribution: K 7; T 2, 3, 6; Z; P; Mozambique, S. Malawi, E.
Zimbabwe. At Kilimanjaro in the area of the Chagga homegardens
widespread.
Pollination: butterflies (Fischer, 2004).
5.2.12. Basellaceae
Short-lived perennials, usually glabrous vines, or decumbent to
procumbent herbs. A tropical and subtropical family of four genera
and about 20 species centred in the New World (Sperling and
Bittrich, 1993).
Pollination: the sweet-smelling flowers of Anredera indicate
insect pollination, but no observations are reported yet, nor about
the pollination of species in the other genera (Sperling and Bittrich,
1993).
Pollen morphology: pollen grains of Anredera, Tournonia, Ullucus
and three species of Basella are spheroidal and pantocolpate or
pantoporate with a spinulose punctuate tectum, other species of
Basella are of cuboidal shape (with a spheroidal nexine layer)
showing a colpus on each of the six faces. An endexine is either
absent or present in the aperture region (Sperling and Bittrich,
1993, and literature herein).
Basella alba L (Plate 8, 22e24).
Shape: quincuadrangular ~33  33 mm.
Apertures: 6-colporate, elliptic pori, narrow colpi of 1/3 length.
Exine: tectate, recticulate, high collumellae.
Habitus: Glabrous annual or shortly lived perennial, succulent
tangled twiner. Stems much-branched, 2e10 m long.
Habitat: In thickets, forest edges, margins of cultivated land and
swampy ground, frequently by rivers or streams; 0 (cultivated)-
2450 m. At Kilimanjaro forest clearings, tall herb vegetation in
montane riverine forest, Chagga homegardens and roadsides, from
970 to 2500 m.
Distribution U2, 4; K1, 3, 4, 6; T1-4, 6, 7; Z; Madeira, West Africa
to Cameroon Republic, S. Tome , DR Congo Republic, Sudan Republic,
Ethiopia, Rwanda Republic, Mozambique, Malawi, Zambia, Angola,
rare in central Africa, Asia to China, Japan, Philippines, Borneo, Fiji
and Hawaii, also in West Indies, Brazil and Guiana, almost certainly
indigenous in Africa. At Kilimanjaro scattered.
Pollination: Not known.
5.2.13. Begoniaceae
Herbs (mostly), or shrubs, or lianas. 920 species in 3e5 genera.
Sub-tropical and tropical. Pantropical, concentrated in America
(Wilde, 2011).
5.2.13.1. Pollination. Insect pollination, though scarcely observed
and recorded, presumably constitutes the major syndrome leading
to fertilization. Pollinating insects are especially bees (Hymenop-
tera, Apoidea) and hoverflies (Diptera, Syrphidae). (Wilde, 2011,
and literature herein; Wyatt and Sazima, 2011).
5.2.13.2. Pollen morphology. Pollen grains 3-colporate, prolate-
perprolate, small. Exine very thin; TEM study reveals the sexine
to be composed of a tectum and irregularly shaped columellae or an
alveolar layer. Pori with granulate membrane, usually slightly
lalongate. Ornamentation is finely striate in Hillebrandia, less
clearly so in Begonia (Erdtman, 1952; Berg, R. G. van den, 1983,
1985).
337
Begonia johnstonii Oliv (Plate 8, 25e26).
Shape: prolate ~9  13 mm, trilobate.
Aperture: 3-colporate, very long (5/6) and narrow colpi, pori
elliptic broad.
Exine: tectate, exine very thin, finely striate but badly visible.
Habitus: herb, possibly perennial, erect, procumbent or strag-
gling, 0.2e0.6 m high or to 0.9 m long.
Habitat: on steep rocks in forest or in moist sites, less often in
forest without any specification; local, rarely locally common;
750e2400 m. At Kilimanjaro mainly in submontane and lower
montane Mitragyna, Newtonia and Afrocrania riverine and gorge
forests, from 1110 to 2340 m.
Distribution: K1, 4, 6, 7; T2, 3, 6, 7; not known elsewhere. At
Kilimanjaro rare.
Pollination: for other Begonia sp. the principal pollinators are
bees of the Apidae and Halictidae (Wyatt and Sazima, 2011).
5.2.14. Bignoniaceae
Usually trees or lianas, sometimes shrubs, rarely herbs, lianas
mostly with unusual vascular structure. A pantropical family with
104 genera and about 860 species, mainly distributed in the Neo-
tropics (Fischer et al., 2004).
5.2.14.1. Pollination. Most Bignoniaceae are bee-pollinated, but
some genera have corollas adapted to bird-pollination, e.g. Pyro-
stegia, Campsis, Dolichandra, Martinella in the New World, and
Spathodea, Lamiodendron, Fernandoa, Tecoma in the Old World
(Fischer et al., 2004).
5.2.14.2. Pollen morphology. Bignoniaceae show high diversity in
pollen features (Buurman, 1977; Gentry and Tomb, 1979; Gentry,
1980). Apertures range from the most frequently occurring, 3-
colp(or)ate type to monocolpate and inaperturate, to 4e5-
colpate, zonocolpate, pericolpate, spiroaperturate and syncolpate.
The exine maybe tectate, semitectate or intectate and varies from
psilate to reticulate, spinulose or aerolate. Mostly monads but also
tetrads or polyads are known.
Jacaranda mimosifolia D. Don (Plate 8, 27e29).
Shape: subprolate 36e42  50 mm, trilobate.
Apertures: 3-colporate, long (5/6), broad and deep colpi, pori
faintly lalongate.
Exine: tectate, psilate.
Habitus: tree up to 15 m or occasionally a shrub 2.5e3 m tall.
Habitat: cultivated. At Kilimanjaro planted as an alley tree, from
900 to 1610 m.
Distribution: native of Argentina and Bolivia. Widely cultivated
in East Africa in gardens and as a street tree. At Kilimanjaro below
the forest belt scattered.
Pollination: insects, primarily by Bombus spp. (Galetto, 1995).
Markhamia lutea (Benth.) K. Schum (Plate 8, 30e31 & Plate 9, 1).
Shape: subprolate, trilobate ~40 mm.
Apertures: 3-colpate, syncolpate, wide and deep colpi, apertural
membrane fissured and granular.
Exine: tectate, microreticulate, homobrochate.
Habitus: tree (5-) 15e21 (24) m tall or sometimes shrubby
1.2e4.5 m tall.
Habitat: wooded grassland, submontane and riverine forest,
evergreen rain forest; 1500e1900 m. At Kilimanjaro in colline-
lower montane Mitragyna, Newtonia and Afrocrania riverine and
gorge forests, as well as in Chagga homegardens, from 780 to
1650 m.
Distribution: U 1e4; K 1, 3e5; T 1, 3, 4 (planted in T2); West
Africa from Ghana to Cameroon, DR Congo-Kinshasa, Rwanda,
Burundi. At Kilimanjaro scattered.
Pollination: pollinated by a wide range of bees including
338 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386

Plate 9. BIGNONIACEAE: 1 Markhamia lutea, 2e3 Tecoma stans, BOMBACACEAE (Malvaceae): 4e5 Adansonia digitata, Boraginaceae: 6e7 Cordia monoica, 8e10 Ehretia bakeri, 11e12
Heliotropium rariflorum ssp. hereroense, 13e14 Lithospermum afromontanum, BURSERACEAE: 15e17 Boswellia neglecta, CAESALPINIACEAE: 18e20 Caesalpinia decapetala, 21e24
Cassia spectabilis, 25e27 Delonix regia, 28e29 Pterolobium stellatum, CAMPANULACEAE: 30e31 Wahlenbergia abyssinica.

carpenter bees, and larger leafcutter bees (Martins, 2014).
Tecoma stans (L.) Juss. ex Kunth (Plate 9, 2e3).
Shape: circular, spheroidal to trilobate ~33 mm.
Apertures: 3-colpate, syncolpate, long, deep and broad colpi
with granular aperture membrane.
Exine: tectate, microreticulate/granulate, sexine thinner around
colpi.
Habitus: shrub or small tree, 2.5e10 m tall.
Habitat: roadsides, Aristida-Harpachne grassland, rocky places,
sandy lake shores; 750e1900 m. At Kilimanjaro planted as orna-
mental shrub, escaped and naturalized in disturbed savanna
woodlands, from 1000 to 1280 m.
Distribution: K 1, 2, 4; T 1, 4, 5; native of America fom Florida to
Argentina, Bolivia and Peru and West Indies, now cultivated widely
and often naturalized. At Kilimanjaro in savanna scattered.
Pollination: bees, wasps and hawkmoths (Jonathan et al., 2009;
Wojcik, 2011).
5.2.15. Bombacaceae ¼ subfamily Bombacoideae (Malvaceae)
Species described here under the former family of Bombacaceae
today belong to the family of Malvaceae. Therefore, detailed in-
formation is given for Malvaceae or the correspondant subfamily
(see also Malvaceae section of this atlas).
The subfamily Bombacoideae is mostly present in the New
World tropics and contains 27 genera (ca. 250 species); a few of
those genera (Adansonia, Bombax, Camptostemon, Lagunaria), con-
taining ca. 19 species, are restricted to the Old World tropics. Spe-
cies are usually trees, often buttressed and/or with a bottle-shaped
trunk, and/or with chunky spines, rarely shrubs or climbers
(Klitgård, 2013).
5.2.15.1. Pollination. Most Bombacoideae have large, massive,
sometimes gigantic flowers usually of the brush type, which are
pollinated by vertebrates, yet occasionally also by bees (Bayer and
Kubitzki, 2003).
5.2.15.2. Pollen morphology. Roughly, the pollen types of the Mal-
vaceae can be divided into a few main types (Bayer and Kubitzki,
2003): the Grewia-type, the Tilia-type, the Helicteres-type, the
Bombax-type and the Malva-type. The Bombax-type which is
 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
mainly found within the Bombacoideae pollen grains are 3-porate
or 3-colporate, peroblate-oblate speroidal (diameter 25e120 mm).
The sexine is often reticulate, sometimes provided with warts or
spinules (Adansonia, Pentaplaris, and some Pachira) (Erdtman,
1952). The enormous variation of shapes, apertures, exine struc-
ture and ornamentation has been described by Nilsson and Robyns
(1986). The species Adansonia digitata, A. kilima, A. gregorii and
A. madagascariensis can be distinguished by the pollen grain size
and the number of spines (Pettigrew et al., 2012).
Adansonia digitata L (Plate 9, 4e5).
Bombax-type.
Shape: circular to subprolate, spheroidal ~58 mm.
Apertures: 3-porate, sub-circular pori Ø 10 mm with annulus and
irregular margins.
Exine: tectate, perforate, verrucate, bacculate with small, blunt
spines with a density of 51e100 spines per 1000 mm2.
Habitus: massive deciduous tree, usually up to 18 m high.
Habitat: coastal bushland, deciduous bushland, woodland,
wooded grassland, semi-desert Chrysopogon vegetation, often left
standing in cultivated areas; 1e1250 m. At Kilimanjaro in savanna
from 800 to 1000 m.
Distribution K 4, 6, 7; T 1e8; Z; P; widespread in the drier areas
of tropical and southern Africa. At Kilimanjaro in savanna rare.
Pollination: fruit bats, lemurs, and long-tongued hawkmoths as
nocturnal pollinators (Baum, 1995).
5.2.16. Boraginaceae ¼ cordiaceae & Ehretiaceae
Trees, or shrubs, or annual to perennial herbs, or lianas (a few).
Holarctic, Palaeotropical, Neotropical, Cape, Australian, and Ant-
arctic. Temperate to tropical. Cosmopolitan, but fewer in cool
temperate and tropical regions, and with a strong Mediterranean
concentration. 2000 species in about 120 genera (Watson and
Dallwitz, 1992 onwards).
5.2.16.1. Pollination. Flower morphology and fragrance suggest
that Cordiaceae are typically zoophilous. However, observations on
pollination have been rarely published. For small, white flowers of
many Ehretiaceae insects like Hymenoptera, Diptera, Lepidoptera,
Coleoptera, or Thysanoptera have been recorded. The large, red
flowers of the subclades Collococcus p.p. and Sebestena p.p. of Cordia
suggest bird pollination. Furthermore, the frequent occurrence of
exserted anthers indicate the possibility of wind pollination
(Gottschling, 2003, and literature herein).
5.2.16.2. Pollen morphology. Pollen grains often 3-colporate (rarely
3-colporidate), suboblate to prolate. Sexine usually as thick as
nexine or thicker (exine stratification often obscure) (Erdtman,
1952).
Cordia monoica Roxb (Plate 9, 6e7).
Shape: spheroidal, circular ~46 mm.
Apertures: 3-colporate, colpi 2/3 in length and narrow, colpi
membrane fissured towards poles, pori elliptic tall but not very well
defined.
Exine: tectate, sexine as thick as nexine, exine microechinate,
spines with blunt tips.
Habitus: much-branched spreading shrub or small tree
branched from the base, 1.5e8 (15) m.
Habitat: very universial, from wet evergreen forest to Acacia
woodland, Acacia-Commiphora bushland and Acacia-Euphorbia
thicket in grassland, coastal thicket, etc. often riverine; 0e1825 m.
At Kilimanjaro in dry savanna forest and woodland; 920e1400 m.
Distribution: U 1e4; K 1e7; T1e3, 5e8; Z; P; Sudan, Ethiopia,
Somalia, also Yemen, Oman, Iran and India. At Kilimanjaro
scattered.
Pollination: not known for this species. Other Cordia sp. are
339
pollinated by birds, bees, moth and flies (e.g. Opler et al., 1975;
McMullen, 2012).
Ehretia bakeri Britten (Plate 9, 8e10).
Shape: prolate ~20  30 mm, triangular to slightly hexalobate.
Apertures: 6-heterocolporate, 3-colpori-3-pseudocolpate, colpi
long (5/6 to syncolpate) and narrow, constricted at the equator,
pseudocolpi shorter, faint and wider, pori elliptic broad but less
distinct than in Cordia.
Exine: tectate, faintly reticulate, exine thicker at poles.
Habitus: shrub or small deciduous tree 1.8e6 m tall.
Habitat: Pterocarpus, Pleurostyla, Brachystegia, Sclerocarya
thicket forest, thicket on shallow soil over rock or on coral rag,
bushland, inner edges of mangrove swamps and gneiss slopes;
0e1050 m. At Kilimanjaro in savanna woodland at 900e1000 m.
Distribution: K 7; T2, 3, 6, 8; P; Kenya (Teita, Kwale, Lamu),
Tanzania (Moshi, Pangani, Morogoro, Pemba), not known else-
where. At Kilimanjaro in savanna scattered.
Pollination: insects (Gottschling, 2003).
Heliotropium rariflorum subsp. hereroense (Schinz) Verdc (Plate
9, 11e12).
Shape: rectangular tall ~23  30 mm, circular/trilobate to slightly
hexalobate.
Apertures: 6-heterocolpate (3-colporate and 3-pseudocolpate)
with narrow colpi 5/6, pseudocolpi shorter than colpi and also
narrow, pori elliptic tall with annulus.
Exine: tectate, sexine slightly thicker than nexine, psilate.
Habitus: much-branched woody subshrub or subshrubby herb
20e80 cm tall. Rootstock ± simple, often very woody, usually
slender than 1 cm.
Habitat: grassland and bushland in dry areas, mainly Com-
bretum-Commiphora-Lannea, Commiphora-Dobera and Commi-
phora-Acacia associations, often the dominant or co-dominant
undershrub; in T6 on sand-banks subject to submergence;
100e1350 m. At Kilimanjaro from 900 to 1000 m in savanna
grassland.
Distribution: K 1, 2, 4, 6, 7; T 2, 3, 6; Kenya (Northern Frontier
Province, Masai, Teita), Tanzania (Masai, Lushoto, Kilosa, Moshi),
Sudan, Ethiopia, Somalia, Angola, Namibia. At Kilimanjaro in
savanna rare.
Pollination: not known for this species. Generally, the flowers of
Heliotropium are adapted to insect pollination (Singh et al., 2009).
Lithospermum afromontanum Weim (Plate 9, 13e14).
Shape: subisopolar-heteropolar, prolate/rectangular tall
~8  15 mm, constricted at the equator, quadrangular.
Apertures: 4-colporate, colpi long (4/5) and narrow, membrane
faintly granular; pori elliptic broad with costae, situated closer to
one pole than to the other.
Exine: tectate, exine very thin <0.5 mm, stratification obscure,
psilate.
Habitus: perennial often straggling subshrubby herb 0.3e1.5 m
tall.
Habitat: montane grassland, clearings in bamboo, bushland and
shrub, Erica and Lycopodium associations, Juniper forest; (1560-)
1800e3950 m. At Kilimanjaro in upper montane Erica excelsa and
Hagenia riverine forest, from 2450 to 2980 m.
Distribution: U 1e3; K 2e6; T 2e4, 7; Uganda (Karamoja, Kigezi,
Mbale), Kenya (Elgeyo, N. Nyeri, Masai), Tanzania (Moshi, Mbulu/
Masai, Mbeya), E. Zaire, Ethiopia, Malawi, Zambia, Zimbabwe, South
Africa. At Kilimanjaro rare.
Pollination: not known for this species. Lithospermum is known
to be pollinated by bees and butterflies (Dobzhansky et al., 1975).
5.2.17. Burseraceae
Trees or shrubs. Approximately 700 species in 19 genera in the
tropics and subtropics, represented by few taxa in some warm
340 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
temperate areas (Daly et al., 2011).
5.2.17.1. Pollination. By insects and specifically with small gener-
alist insect species (Daly et al., 2011, and literature herein).
5.2.17.2. Pollen morphology. Pollen grains usually 3-colporate,
usually angulaperturate, occasionally fossaperturate. Pollen shape
either suboblate oblate-spheroidal to prolate-spheroidal sub-
prolate to prolate, in polar view circular, rounded-triangular,
triangular, and triangular-lobed. The colpi may be very short or
long. Endoapertures usually lalongate or per-lalongate, circular,
subcircular or, rarely, lolongate. Exine tectate or semitectate and
either psilate, psilate-perforate, foveolate, low relief rugulate,
scabrate-rugulate, perforate-rugulate, striate, striate-perforate,
striate-reticulate (Daly et al., 2011).
Boswellia neglecta S. Moore (Plate 9, 15e17).
Shape: circular, subprolate to spheroidal, ~48  54 mm.
Apertures: 3-colporate, brevicolpate, colpi narrow, pori elliptic
broad ~7  10 mm.
Exine: tectate, sexine about as thick as nexine, psilate, scabrate.
Habitus: shrub or small tree up to 8 m tall.
Habitat: Acacia, Commiphora bushland on basement complex or
lava and red sandy soil; 200e1350 m; rainfall 250e600 mm. At
Kilimanjaro in savanna woodlands from 920 to 1100 m.
Distribution: U 1, K 1e4, 7; T 2, 3; E. Ethiopia, N., C. and S. So-
malia. At Kilimanjaro in savanna scattered.
Pollination: bees (Wigrup, 2005; Ramawat, 2010).
5.2.18. Caesalpiniaceae
Trees, shrubs, perennial herbs or lianas. The family is distributed
in the Paleotropics, Neotropics, the Cape Region, and Australia. A
family of 150 genera, 2200 species; widespread in tropical and
subtropical regions (Brenan, 2006).
5.2.18.1. Pollination. Usually insects; pollination mechanism can be
conspicuously specialized, or unspecialized (Watson and Dallwitz,
1992 onwards).
5.2.18.2. Pollen morphology. The basic pollen type in the family is a
3-colporate monad with a perforate or finely reticulate tectum
(Guinet and Ferguson, 1989). Grains are shed in monads and very
rarely aggregated (Afzelia, Diptychandra); when aggregated, in
tetrads.
Caesalpinia decapetala (Roth) Alston (Plate 9, 18e20).
Shape: circular, spheroidal ~54 mm.
Apertures: 3-colporate, colpi very broad and long (5/6) with
granular membraned margo, colpi tapering at poles, colpoid streaks
(intectate) very distinct, pori elliptic tall.
Exine: intectate, in the colpus area bordered by tectate meso-
colpi which are coarsely reticulate, heterobrochate, sexine thicker
than nexine.
Habitus: climbing or straggling bushy shrub 2.5e10 m high.
Habitat: clearings in lowland rain-forest, scattered tree grass-
land, bushland; 880e2130 m. At Kilimanjaro in disturbed riverine
forest, thicked edges, hedges, plot boundaries in Chagga home-
gardens, waste land, from 900 to 1530 m.
Distribution: U 2, 4; K ?3, 4, 5; T 1e3, 6, 7; originally from
tropical and subtropical Asia, but now widely cultivated and often
naturalized; in Africa from the Flora area southwards to Angola and
South Africa. At Kilimanjaro below the forest belt widespread.
Pollination: by various insect species (Byrne et al., 2011).
Cassia spectabilis DC (Plate 9, 21e24).
Shape: subprolate ~23  28 mm, trilobate-circular ~28 mm.
Apertures: 3-colporate, long colpi (4/5) constricted at equator
with blunt ends towards the poles, pori indistinct and circular.
Exine: tectate, sexine thinner than nexine, psilate.
Habitus: small, rounded deciduous tree, 7e10 m (max. 15) tall.
Habitat: exotic species which is a common garden plant that
also invades; forest margins, savanna, riverbanks, roadsides, waste
ground and plantations. At Kilimanjaro planted as alley tree, from
820 to 1380 m.
Distribution: native of tropical America. At Kilimanjaro
scattered.
Pollination: insects, predominantly by bees (Manente-Balestieri
and Machado, 1999).
Delonix regia (Bojer ex Hook.) Raf (Plate 9, 25e27).
Shape: circular-subprolate, trilobate-spheroidal ~56 mm.
Apertures: 3-colporate, broad and short (1/2) colpi with gran-
ular margins, pori badly definded.
Exine: tectate, sexine thicker than nexine, macroreticulate,
heterobrochate.
Habitus: tree with 10e15 m of height. As the trees mature, they
develop broad umbrella-shaped crowns, and are often planted for
their shade-giving properties.
Habitat: found in Madagascar's dry deciduous forests. In the
wild it is endangered, but it is widely cultivated elsewhere. At
Kilimanjaro planted as an ornamental tree from 800 to 1000 m.
Distribution: very rare native of Madagascar, cultivated in East
Africa as throughout the tropics. At Kilimanjaro scattered.
Pollination: ornithophilous, birds and butterflies, classic orni-
thophilous species of the Old Worlds that acquired ornithophily
through butterfly-pollinated intermediaries (Endress, 1996, and
literature herein).
Pterolobium stellatum (Forssk.) Brenan (Plate 9, 28e29).
Shape: circular-triangular~27 mm, spheroidal-subprolate.
Apertures: 3-colporate, long (5/6) and very broad colpi with
granular membraned margo, small lolongate pori with vestibulum.
Exine: tectate, sexine slightly thicker than nexine, reticulate.
Habitus: scrambling or climbing, rarely semi-erect shrub
2e15 m high.
Habitat: upland dry evergreen forest (? also on edges of upland
rain-forest), riverine forest, deciduous woodland, sometimes on
termite-mounds in grassland; 850e2290 m. At Kilimanjaro mainly
in colline-submontane forests (dry savanna forest, riverine forests,
Croton-Calodendrum forest), from 780 to 1900 m.
Distribution: U 1e4; K 1e6; T 1e7; widespread in eastern Africa,
from Eritrea and the Sudan Republic southwards to the Transvaal,
also in Arabia. At Kilimanjaro in lower located forests widespread.
Pollination: bee-pollinated (Wubie et al., 2014).
5.2.19. Campanulaceae
Herbaceous perennials, less often annuals or biennials, herba-
ceous or woody lianas (sometimes twining), pachycaul rosette
plants, subshrubs, shrubs, treelets, or trees to 15 m tall, typically
terrestrial, rarely aquatic or epiphytic. 84 genera and almost 2400
species. The family is cosmopolitan, with representatives on six
continents and several of the world's archipelagos (Lammers,
2009).
5.2.19.1. Pollination. Entomophilous; mechanism conspicuously
specialized, including bees, flies, wasps, butterflies, and settling
moths. Species with zygomorphic corollas have a more restricted
range of specialized, not infrequently vertebrate vectors (Lammers,
2009, and literature herein).
5.2.19.2. Pollen morphology. Pollen grains are spheroidal or vari-
ously compressed (oblate or prolate). Grains colpate, colporidate,
colporate or porate. In most genera, grains are 3e4(-5)-porate.
Campanuloideae show greater diversity. Surface ornamentation is
variable. Pollen grains are dispersed as monads (Erdtman, 1952;
 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386 341

Dunbar, 1975). Angola, Zaire, Zambia, Malawi, Mozambique, Zimbabwe, South

Wahlenbergia abyssinica (Hochst. ex A. Rich.) Thulin (Plate 9,
30e31 & Plate 10, 1).
Shape: circular, spheroidal ~30 mm.
Apertures: 3-porate, circular pori Ø ~6 mm sunken with narrow
annulus.
Exine: tectate, sexine thinner than nexine, microechinate.
Habitus: perennial or annual herb from a taproot, which
becomes ± long and woody in old perennating specimens. Stems
few to many, erect or rarely ± decumbent, 0.12e0.90 m tall.
Habitat: woodland (usually deciduous) or grassland, forest
clearings, old cultivations, roadsides, usually in sandy or rocky soils;
0e2700 m. At Kilimanjaro mainly in anthropogenic dry grassland of
the submontane zone with Hyparrhenia rufa, Satureia abyssinica,
Aristida adoensis and Crepis carbonaria, also in dry lower montane
Bulbostylis meadows, from 1130 to 2130 m.
Distribution: K 1e4, 6, 7; T 2, 3, 5e8; Z; Ethiopia, Somalia,
Africa, Madagascar. At Kilimanjaro in the submontane cultivation
belt scattered.
Pollination: insect; no further details known.
5.2.20. Capparaceae
Annual or perennial herbs, subshrubs, shrubs or rarely trees,
sometimes climbing, scrambling or rarely lianas. A family of 29
genera and about 700 species of worldwide distribution, prefer-
entially in tropical and subtropical regions with pronounced sea-
sonal drought, including regions with Mediterranean climate (Kers,
2003).
5.2.20.1. Pollination. Different pollinating agents; in the large
genera Cleome and Capparis bees, hummingbirds, hawkmoths, and
bats are involved in pollination. Protandry and self-compatibility
are common, self-incompatibility has been reported (Kers, 2003,

Plate 10. CAMPANULACEAE: 1 Wahlenbergia abyssinica, CAPPARACEAE: 2e3 Cleome hirta, 4e6 Maerua grantii, CARYOPHYLLACEAE: 7e8 Cerastium indicum, 9e10 Uebelinia
rotundifolia, Celastraceae: 11e13 Hippocratea africana, 14e16 Maytenus acuminata, CHENOPODIACEAE: 17e18 Chenopodium ambrosioides, COMBRETACEAE: 19e21 Combretum
apiculatum, 22e24 Combretum psidioides subsp. psidioides, COMMELINACEAE: 25e26 Aneilema aequinoctiale, 27e28 Aneilema johnstonii, 29e30 Commelina foliacea, CONNARACEAE:
31e33 Rourea thomsonii, CONVOLVULACEAE: 34e35 Astripomoea hyoscyamoides subsp. hyoscyamoides, 36e38 Convolvulus kilimandschari, CRASSULACEAE: 39e42 Crassula
alsinoides.
342 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
and literature herein).
5.2.20.2. Pollen morphology. Capparaceae are stenopalynous. Pol-
len grains (2-) 3 (4)-colporate (-colporidate), usually subprolate
or prolate, in polar view circular or rarely subtriangular. Sexine as
thick as nexine or thicker, tectate (sometime perforate). Some
species are microechinate (Erdtman, 1952). The apertures are
compound, and the ectoapterure extends to near the poles. The pori
are circular, lalongate or lolongate, and the operculum is diffuse or
distinct in the colporate type (Kers, 2003; Kubitzki, 2003a).
Cleome hirta (Klotzsch) Oliv (Plate 10, 2e3) (since 2008 in
Cleomaceae (APG, 2009)).
Shape: prolate ~10  14 mm, trilobate ~10 mm.
Apertures: 3-colporate, long (5/6) and narrow colpi, pori
circular.
Exine: tectate, sexine as thick as nexine, microreticulate-
rugulate.
Habitus: annual or short-lived perennial herb. Well branched,
up to 1.5 m tall.
Habitat: deciduous bushland and grassland, becoming a weed of
roadsides and cultivated ground; 0e1800 m. At Kilimanjaro on
roadsides, fallow arable fields and waste places in the savanna, from
850 to 1190 m.
Distribution: U 1e4; K 1, 2, 4e7; T 1e8; Mozambique, Nyasa-
land, N. and S. Zimbabwe, Rwanda and DR Congo Republics, also
Ethiopia. At Kilimanjaro in the savanna scattered.
Pollination: bees, butterflies, and hawkmoths (Martins, 2014).
Maerua grantii Oliv (Plate 10, 4e6).
Shape: (sub)prolate ~22  30 mm, trilobate.
Apertures: 3-colporate, colpi long (5/6), wide at equator and
tapering at poles, large pori elliptic broad/lalongate.
Exine: tectate, sexine noticeably thicker than nexine, reticulate,
homobrochate.
Habitus: small twiggy shrub, much branched and somewhat
straggeling, up to 2.5 m tall or e after burning e flowering on erect
branched almost herbaceous shoots 0.6 m tall.
Habitat: deciduous woodland, bushland, grassland with scat-
tered trees and riverine forest; persisting on roadsides and waste
places; 50e1050 m. At Kilimanjaro in savanna grasslands, from
1000 to 1100 m.
Distribution: K 1, 7; T 2e6, 8; Mozambique. At Kilimanjaro rare.
Pollination: various insects (hawkmoths, and carpenter -, honey
- and solitary bees) (Martins, 2014).
5.2.21. Caryophyllaceae
Annual or perennial herbs, or subshrubs, rarely shrubs or small
trees. About 86 genera with ca. 2200 species, mainly distributed in
the temperate regions of the northern hemisphere with a center of
the Mediterranean and Irano-Turanean regions (Bittrich, 1993).
5.2.21.1. Pollination. Most species of Caryophyllaceae are ento-
mophilous. Wind pollination is not reported with certainty
(Bittrich, 1993).
5.2.21.2. Pollen morphology. Pollen grains are suboblate-subprolate
(if 3-colpate), and spherical or ± rounded polyhedral (if porate or
pantoporate). The exine is tectate, the tectum is mostly perforate or
occasionally anulopunctate, or rarely reticulate, and finely spinu-
lose. The endexine is thick, thin, or even absent. The tectum is
rather thick and separated by more or less numerous columellae
from the foot layer, sometimes columellae are hanging. Pollen
grains are mostly (3) colpate, or rarely porate (pantoporate)
(Erdtman, 1952; Bittrich, 1993, and literature herein).
Cerastium indicum Wight & Arn (Plate 10, 7e8).
Shape: circular, spheroidal ~30 mm.
Apertures: periporate, 9e15 circular pori Ø ~5 mm with annulus.
Exine: tectate, microreticulate, columellate, more microbrochi
than columellae, heterobrochate, lumina smaller around pori.
Habitus: perennial herb, with spreading and ascending slender
branches, stems 20e60 cm long.
Habitat: grassland, bushland, forest margins and glades, road-
sides, pathsides, weed of cultivated land; 1050e2900 m. At Kili-
manjaro mainly in upper montane Erica excelsa forest, subalpine
Erica bushland, also in wet meadows of the lower montane zone,
from 1610 to 2840 m.
Distribution: U 2, 3; K 3e7; T 2, 3, 6, 7; A.-E. Sudan. At Kili-
manjaro rare.
Pollination: insects (e.g. Willemstein, 1987).
Uebelinia rotundifolia Oliver (Plate 10, 9e10).
Shape: circular, spheroidal ~33 mm.
Apertures: periporate, pores ~3 mm with thin annulus.
Exine: tectate, microreticulate, columellate, more microbrochi
than columellae, heterobrochate, lumen smaller around pori.
Habitus: procumbent herb with branching often elongated
stems rooting at many of the nodes.
Habitat: forest and wet shady stream banks, 2600e2900 m. At
Kilimanjaro in upper montane Hagenia and Erica excelsa forests,
from 2950 to 3200 m.
Distribution: T2; endemic to Kilimanjaro, very rare.
Pollination: probably insects.
5.2.22. Celastraceae
Erect or scandent trees, shrubs, lianas, or annual or perennial
herbs, rhizomatous shrubs, ericoid subshrubs, or epiphytic shrubs.
A subcosmopolitan family of 98 genera and about 1211 species that
is most diverse in the tropics and subtropics, with fewer temperate
species (Simmons, 2004).
5.2.22.1. Pollination. Pollinated by bees, beetles, flies, wasps, and
even ants (Simmons, 2004, and literature herein).
5.2.22.2. Pollen morphology. Pollen grains usually 3-colporate,
suboblate-subprolate. Sexine thicker than nexine, usually reticu-
late, simplibacculate. Size of lumina decreases towards colpi
(Erdtman, 1952). The most thorough survey of pollen in Celas-
traceae was completed by Lobreau-Callen (1977). Within the
Celastroideae, pollen is isopolar and shed in monads, whereas in
Hippocrateoideae, Salacioideae and Lophopetalum, pollen may also
be shed in tetrads or polyads of eight or 16 grains. The apertures of
grains shed in tetrads or polyads may be irregularly positioned.
Grains are generally 3-colporate, but bicolporate and tetracolporate
grains have been reported. Some genera in which grains are shed in
tetrads or polyads have triporate grains. Grains are suboblate to
subprolate, small to medium sized, with reticulate (rarely croto-
noid) sculpturing (Lobreau-Callen, 1977).
Hippocratea africana (Willd.) Loes. ex Engl (Plate 10, 11e13).
Shape: suboblate, triangular convex ~26 mm.
Apertures: 3-colporate, circular-squared pori with annulus,
colpi long (4/5) and narrow with margo.
Exine: tectate, sexine thicker than nexine but thinner towards
apertures, visible collumella, reticulate, lumina size decreases to-
wards colpi.
Habitus: liana.
Habitat: evergreen forest, fringing forest and thickets;
500e1675 m. At Kilimanjaro in colline-lower montane riverine and
gorge forests, Croton-Calodendrum and Cassipourea forests, from
980 to 2100 m.
Distribution: U2, 4; K4e6; T3; widely distributed in Africa from
Mauritania east to Ethiopia and south to South Africa (Transvaal)
and Botswana, India, Sri Lanka, Burma, Laos. At Kilimanjaro in the
 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
lower located forests scattered.
Pollination: insects (e.g. Simmons, 2004).
Maytenus acuminata (L.f.) Loes (Plate 10, 14e16).
Shape: trilobate, circular ~30 mm.
Apertures: 3-colporate, square pori, colpi (2/3) narrow and
tapering towards poles.
Exine: tectate, sexine considerably thicker than nexine, retipi-
late/reticulate.
Habitus: shrub or tree (0.3-) 1e20 m high, erect or ± spreading.
Habitat: understorey and margin of upland or riverine forest;
1050e3300 m. At Kilimanjaro widespread in lower-upper montane
(riverine, Cassipourea, Ocotea, Podocarpus and Juniperus) forests,
from 1740 to 2990 m.
Distribution: U 2, 4; K 4, 7; T 1e4, 6, 7; Zaire (Kivu), Rwanda and
southward to South Africa (Cape). At Kilimanjaro in the forest
widespread, but in low frequencies.
Pollination: insects (e.g. Willemstein, 1987).
5.2.23. Chenopodiaceae
Annual or perennial, often halophytic herbs, sometimes shrubs
or small trees. A family of the temperate and subtropical regions
with ca. 100 genera and 1400 species; many species are dominant
constituants of both salt-marshes, deserts and semi-deserts (Kühn,
1993).
5.2.23.1. Pollination. Formerly the Chenopodiaceae were supposed
to be uniformly wind-pollinated. In some genera (mostly of the
Salsoloideae) entomophily plays a role. Self-pollination can also be
found (Kühn, 1993).
5.2.23.2. Pollen morphology. Chenopodiaceae are stenopalynous in
having only pantoporate pollen. The grains have an exine covered
by a thick, minutely perforate tectum with short spinules. Sexine
usually considerably thicker than nexine, tectate. The pori, which
occur in varying numbers, consist of reduced pointed flecks of exine
underlain by lamellar plates, two pollen types are distinguished by
Nowicke and Skvarla (1979): one spheroidal, with numerous small
pori, the other polyhedral, with fewer, larger and slightly sunken
pori (Kühn, 1993).
Chenopodium ambrosioides L (Plate 10, 17e18).
Shape: circular, spheroidal ~23 mm.
Apertures: periporate, many (>40) small circular pori Ø ~1 mm
with narrow annulus.
Exine: tectate, collumellate, verrucate.
Habitus: herb up to 1.2 m high, usually annual, rarely a short-
lived perennial, polymorphic.
Habitat: weed near habitations, sometimes on permanent ways.
Certainly not native; altitudinal range incompletely known, but
recorded at about 1220 m. At Kilimanjaro in ruderal vegetation in
Chagga homegardens, on roadsides, fallow arable fields and waste
places in the savanna and submontane zone, from 960 to 1430 m.
Distribution: U 4; K 4; T 1, 4, 6; throughout the tropical and
subtropical regions of the world, especially polymorphic in Amer-
ica. At Kilimanjaro rare.
Pollination: mainly wind-pollinated (Coile and Artaud, 1997).
5.2.24. Combretaceae
Trees, shrubs, subshrubs or lianas, sometimes mangroves, rarely
spiny. A pantropical family with 14 genera and ca. 500 species
(Stace, 2010).
5.2.24.1. Pollination. Presumably pollinated by sunbirds, although
no direct observations have been made; a wide range of insects,
including beetles, flies, bees and butterflies; some species are
pollinated by hummingbirds but other birds, butterflies and
343
monkeys also play their part (Stace, 2010, and literature herein).
5.2.24.2. Pollen morphology. Pollen grains 3-colporate, prolate
spheroidal-prolate. Sexine evidently thinner than nexine,
reticulate-polybrochate or without distinct pattern (Erdtman,
1952).
Combretum apiculatum Vent (Plate 10, 19e21).
Shape: subprolate ~23  30 mm, trilobate-hexalobate.
Apertures: 6-heterocolporate, 3-colpori-3-pseudocolporate,
colpi long (5/6) and narrow, pseudocolpi shorter (2/3),
pori ± circular Ø ~4.5 mm.
Exine: tectate, psilate-scabrate.
Habitus: small tree up to 10 m high, rarely a shrub.
Habitat: Brachystegia woodland, wooded grassland and Acacia,
Commiphora bushland, common on rocky hill slopes; 70e1800 m.
At Kilimanjaro in savanna woodland, dry and riverine forests, from
1000 to 1100 m.
Distribution: K 1, 4, 7; T 1e8; Botswana, Zambia, Zimbabwe,
Malawi, Mozambique, Angola, South and South West Africa. At
Kilimanjaro in the savanna rare.
Pollination: probably different pollinators like insects, hum-
mingbird, bats (Schemske, 1980; Bernardello, 1994; Ekeke and
Agbagwa, 2015).
Combretum psidioides Welw. subsp. psidioides (Plate 10, 22e24).
Shape: rectangular tall to prolate ~25  34 mm, trilobate-
hexalobate ~28 mm.
Apertures: 6-heterocolporate, 3-colpori-3-pseudocolporate,
colpi long (5/6), narrow and constricted at equator with thin margo,
pseudocolpi shorter (2/3) and slightly wider, pori elliptic/rectan-
gular tall.
Exine: tectate, scabrate/granulate.
Habitus: tree up to 17 m high or large bush.
Habitat: Brachystegia and other deciduous woodlands, often on
rocky hill slopes; 900e1800 m.
Distribution: K 4; T 1, 3, 4, 7, 8; Zaire, Malawi, Mozambique,
Zambia, Zimbabwe, Botswana, Angola, South West Africa.
Pollination: not known for this species. Probably insects like
most other Combretum species.
5.2.25. Commelinaceae
Perennial or occasionally annual, terrestrial, or rarely epiphytic,
or small to large herbs. A mainly tropical and warm temperate
family of about 41 genera and 650 species (Faden, 1998).
5.2.25.1. Pollination. Flowers are chiefly entomophilous and usu-
ally attract a variety of bees and Diptera (Faden, 1998).
5.2.25.2. Pollen morphology. Pollen grains are sulcate except in a
few species of Tinantia, Tradescantia, and Tripogandra (Poole and
Hunt, 1980). Although a number of exine patterns were recog-
nized by Poole and Hunt (1980), Faden and Hunt (Faden and Hunt,
1991) combined them into 2 general types, spinulose and lacking
spines. The pollen grains may be dimorphic in different anthers of
the same flower (Faden, 1998). The exine is thin; sexine usually
thicker than nexine (Erdtman, 1952).
Aneilema aequinoctiale (P. Beauv.) G. Don (Plate 10, 25e26).
Shape: heteropolar, spheroidal, circular/elliptic ~50 mm.
Apertures: monosulcate, sulcus very broad and long (up to 5/6),
aperture margins irregular.
Exine: tectate, exine thin, loosly bacculate.
Habitus: perennial to 2 (3) m long. Roots thin, shoots scram-
bling or decumbent.
Habitat: moist or dry evergreen forest, mist and riparian forest,
especially forest edges and glades, often in dense undergrowth,
along rivers and streams and rarely lakes, Brachystegia woodland,
344 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
thickets and thicket edges, rocky slopes, roadsides and roadside
banks and other moist, disturbed place; 0e1950 m. At Kilimanjaro
in riverine forests as well as in ruderal vegetation in Chagga
homegardens, on roadsides, fallow arable fields and waste places of
the savanna-lower montane zone, from 780 to 2470 m.
Distribution: U 2, 4; K 1, 4, 7; T 1e8; Guinea, Liberia, Ivory Coast,
Ghana, Nigeria, Cameroon, Central African Republic, Equatorial
Guinea, DR Congo, Rwanda, Burundi, Sudan, Ethiopia, Angola,
Malawi, Mozambique, Zimbabwe, Zwaziland, South Africa. At Kili-
manjaro below the forest belt scattered.
Pollination: various insects (flies and bees) are pollinating
agents in this genus (e.g. Faden, 1992; Williams and Adam, 2010).
Aneilema johnstonii K. Schum (Plate 10, 27e28).
Shape: prolate ~31  48 mm, circular/elliptic.
Apertures: monocolpate, sulcus narrow to broad and 2/3 to 4/5
long, aperture margins irregular and fissured.
Exine: tectate, exine thin, wall often slightly thinner on poles,
microbacculate/microverrucate (bacculae/verrucae denser than in
A. aequinoctiale).
Habitus: perennial geophyte, roots thin, shoots tufted,
25e100 cm tall, usually erect to ascending, rarely straggling.
Habitat: Brachystegia (miombo) woodland, wooded grassland,
grassland, Acacia-Commiphora woodland, scrub or thorn bush, in-
selbergs and other rocky habitats, termite hills, edge of mbugas and
flooded depressions, riverine thicket; sandy or clayey soils; light
shade to full sun; 200e2100(-3200) m. At Kilimanjaro in savanna
grassland from 1000 to 1100 m.
Distribution: K 1, 4, 6, 7; T 2e8; Ethiopia, Mozambique, Zambia,
Malawi, Zimbabwe, Botswana. At Kilimanjaro very rare.
Pollination: various insects (flies and bees) are pollinating
agents in this genus (e.g. Faden, 1992; Williams and Adam, 2010).
Commelina foliacea Chiov (Plate 10, 29e30).
Shape: prolate ~18  28 mm, circular/elliptic.
Apertures: monocolpate, sulcus broad, straight and long (4/5),
aperture margins irregular and fissured.
Exine: tecate, thin exine, microverrucate.
Habitus: rhizomatous perennial, shoots erect or ascending or
occasionally trailing, to ±0.3 m high or 1 m long.
Habitat: exceedingly varied: montane moist evergreen forest,
forest paths, edges, and light gaps, dry evergreen forest, lowland
wet evergreen forest, mist forest, Acacia xanthophloea woodland,
wooded grassland, seasonally wet grassland, thickets, montane
scrub, Acacia-Commiphora bushland, roadsides, edges of cultiva-
tion, often in riparian habitats; near sea level to 2300 m. At Kili-
manjaro mainly in ruderal vegetation in Chagga homegardens, on
roadsides, fallow arable fields and waste places, from 800 to
1870 m.
Distribution: U 2; K 1, 2, 4e7; T 2, 3; S Ethiopia, DR Congo-
Kinshasa, Malawi, Zambia. At Kilimanjaro below the forest belt
scattered.
Pollination: generally, insects in this genus, but self-pollination
has also been observed in some species (Oziegbe et al., 2013).
5.2.26. Connaraceae
Small trees, shrubs, or lianas, evergreen or sometimes decidu-
ous. A family comprising 12 genera and 110e200 species, almost
exclusively found in the Tropics. The family is largely restricted to
lowland rainforest, but some species occur in mountains and others
in thickets in savannas (Lemmens et al., 2009).
5.2.26.1. Pollination. Observations on pollination are very scarce.
Many species are sweet scented and probably pollinated by insects.
Bees have been observed visiting flowers (Lemmens et al., 2009).
5.2.26.2. Pollen morphology. Two different pollen types are
encountered. The pollen grains in Jollydoroideae are 4-colpate,
quadrangular, bilateral and subisopolar (bean shaped: faintly
convex-concave). Sexine about as thick as nexine or slightly
thinner, finely reticulate (muri dupli-multibaculate). In Con-
naroideae, pollen grains are 3-aperturate (colpate, porate or col-
porate), suboblate-subprolate. Sexine as thick as nexine or slightly
thicker or thinner, coarsely reticulate (Erdtman, 1952).
Rourea thomsonii (Bak.) Jongkind (Plate 10, 31e33) syn: Jaundea
pinnata (Beauv.) Schellenb.
Shape: spheroidal, circular to trilobate ~21 mm.
Apertures: 3-colporate, porous region indistinct, broad colpi (5/
6) with thin margo tapering towards poles.
Exine: tectate, sexine slightly thicker than nexine but thinning
towards apertures, reticulate, homobrochate.
Habitus: shrub, small tree or liana, up to 25 m.
Habitat: typically, a liana of lowland and upland rainforest, but
occurs as shrub or small tree in forest remnants and cleared agri-
cultural areas, 0e2500 m. At Kilimanjaro mainly in lower-middle
montane riverine, gorge, Cassipourea and Ocotea forests, from
1270 to 2330 m.
Distribution: U 2, 4; K 4, 7; T 1e3, 6, 7; extends from French
Guinea in the west, to the A.-E. Sudan, and southwards through
northern Zimbabwe to Angola. At Kilimanjaro in the lower forest
belt widespread.
Pollination: probably insects (Lemmens et al., 2009).
5.2.27. Convolvulaceae
The Convolvulaceae are a family of herbaceous, twining, or
woody, climbing or trailing vines, shrubs or trees. It is almost
cosmopolitan in distribution, but primarily tropical, with many
genera endemic to individual continents (Austin, 1998).
5.2.27.1. Pollination. Convolvulaceae contain species adapted to
several pollinators such as bees (most frequent), moths, nectar-
feeding birds and bats.
5.2.27.2. Pollen morphology. Pollen grains colpate (sometimes col-
porate?), porate or loxoaperturate. Longest axis from 20 to 210 mm.
Two types: Type 1) Ipomoea-type: grains polyporate, very large
(diameter 90e210 mm), thick exine, echinate. The exine consists of
more or less rodlike elements. The rods often coalesce with the
echinae to form basal rootlets. The length of the echinae varies from
7 mm to 12 mm. Type 2) Other types: grains 20e90 mm, tectum as a
rule distinctly perforate, neither with a thick exine nor echinate (or
at least not with long echinae) (Erdtman, 1952).
Astripomoea hyoscyamoides (Vatke) Verdc. subsp. hyoscyamoides
(Plate 10, 34e35).
Ipomoea-type.
Shape: circular, spheroidal ~71 mm.
Apertures: periporate, many circular pori Ø ~ 2 mm.
Exine: tectate, echinate, spines ~5 mm long with ± blunt tip and
rather narrow base.
Habitus: shrubby short-lived perennial or annual. Stems
0.5e2.4 m tall.
Habitat: Acacia desert-grassland, deciduous bushland, road-
sides, cultivated and other disturbed ground; 50e1200 m. At Kili-
manjaro in ruderal vegetation on roadsides, fallow arable fields and
waste places in the savanna from 990 to 1170 m.
Distribution: K 4, 6, 7; T 1e3, 5e7; Z; not known elsewhere. At
Kilimanjaro in the savanna scattered.
Pollination: not known.
Convolvulus kilimandschari Engl (Plate 10, 36e38).
Other type.
Shape: subprolate ~69  78 mm, circular ~72 mm, grains mostly
torn open along colpi.
 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
Apertures: 3-colporate, pori lolongate and sunken, colpi long (4/
5), recessed with irregular margins.
Exine: tectate, exine very thick, sexine thicker than nexine,
reticulate/retipilate.
Habitus: perennial climber. Stems twining, rarely prostrate, up
to 2 m long.
Habitat: edge of upland rain-forest, moist bamboo thicket and
upland moor, more rarely in upland grassland and secondary
bushland; 1800e3750 m. At Kilimanjaro mainly in upper montane
Podocarpus and Erica excelsa forest, forest edges and clearings, from
1850 to 2900 m.
Distribution: U 3; K 3, 4, 6; T 2; Ethiopia. At Kilimanjaro in the
forest belt rare.
Pollination: it seems, that species in the genus Convolvulus are
mainly pollinated by bees, moth and butterflies (e.g. Prokop and
Neupauerova , 2014).
5.2.28. Crassulaceae
Perennial or rarely annual or hapaxanthic herbs to (sub)shrubs,
rarely aquatics, treelike, epiphytic or scandent, with ± succulent
leaves, sometimes with succulent steems, rhizomes, underground
caudices or succulent roots. A family of 34 genera with ca. 1410
species distributed worldwide, usually in arid and/or rocky habi-
tats, with centers of diversity in Mexico and South Africa (Thiede
and Eggli, 2010).
5.2.28.1. Pollination. Crassulaceae appear to be usually self-
incompatible but Sedum sect. Gormania shows self-compatibility
in varying degrees (Denton, 1979): Five major pollination syn-
dromes are found (Vogel, 1954): Mellitophily and Psychophily are
most common; the sphingophilous syndrome appears to be
restricted to a few Crassula and some Kalanchoe, and thus to Africa.
Ornithophilous flowers are found in species of Kalanchoe, Coty-
ledon, Tylecodon, Echeveria and Dudleya (Parra et al., 1993).
Myophily is assumed for Monanthes.
5.2.28.2. Pollen morphology. Pollen grain usually 3-colporate and
subspheroidal to prolate in equatorial view, ± convex-triangular in
polar view, and 13e38 mm long. Apertures are lalongate. The sexine
is about as thick as the nexine. The tectum is complete and usually
striate, reticulate, rugulate or cerebroid. The striae have a straight or
rarely irregular margin (Monanthes). Colpi are tenuimarginate, with
the thin exine usually protruding at the equatorial part of the colpi
(Erdtman, 1952). Pollen morphology may vary within the same
inflorescence and is thus of restricted systematic applicability
(Thiede and Eggli, 2010).
Crassula alsinoides (Hook. f.) Engl (Plate 10, 39e42).
Shape: (sub) prolate, trilobate, ~16  23 mm.
Apertures: 3-colporate, colpi long (4/5) and narrow, pori circular
to elliptic broad.
Exine: tectate, psilate, exine protruding at equatorial part of
colpi.
Habitus: creeping mat-forming perennial herb. Stems 4-angled,
prostrate or ascending, up to 0.5 m or more long, sparingly
branched.
Habitat: moist ground by streams and swamps in upland forest
and bushland, cultivated land and roadsides; (1300-)
2500e3500 m. At Kilimanjaro in ruderal vegetation in Chagga
homegardens, on roadsides, trampled ground on montane forest
tracks, wet meadows and streamsides, from 1080 to 3080 m.
Distribution: U 2, 3; K 1, 3e5, 7; T 2, 3, 6e8; Fernando-Po,
Cameroon, Zaire, Burundi, Rwanda, Sudan, Ethiopia, Somalia,
Zimbabwe, Malawi, Mozambique, South Africa, Madagascar,
Yemen. At Kilimanjaro scattered.
Pollination: not known.
345
5.2.29. Cucurbitaceae
Climbers or trailers, rarely without tendrils, herbaceous annual
vines or woody perennial lianas, exceptionally trees (Dendrosicyos),
often with tuberous roots or rootstocks or with leafless
and ± succulent stems. The family contains 97 genera and 940e980
species. Its distribution is essentially tropical and subtropical, with
relatively few species reaching the temperate regions of the world
(Schaefer and Renner, 2011).
5.2.29.1. Pollination. Pollen-foraging bees are the predominant
pollinators of Cucurbitaceae. About 86 species of Momordica,
Thladiantha, and a few other genera have oil-secreting trichomes on
the petal bases, and are pollinated by specialized oilbees of the
genus Ctenoplectra (Schaefer and Renner, 2011, and literature
herein).
5.2.29.2. Pollen morphology. An eurypalynous family. Pollen grains
are tectate to intectate, and grains are shed as monads, rarely as
tetrads (Borneosicyos, Gurania, Psiguria); colpate, colporate or
porate, varying in shape from oblate to prolate and in size from
18 mm to 180 mm. Sexine thicker than nexine, reticulate with sim-
plibaculate muri (Erdtman, 1952; Schaefer and Renner, 2011).
Diplocyclos schliebenii (Harms) C. Jeffrey (Plate 11, 1e3).
Shape: spheroidal, circular ~95 mm, shape varies due to thin
exine.
Apertures: 3-porate, large ± circular pori.
Exine: semi (tectate), loosly echinate, distinctly foveolate.
Habitus: rampant climber forming large masses on shrubs at
forest edges and in undergrowth. Stems herbaceous.
Habitat: gaps and margins in upland rain-forest; 1250e1980 m.
At Kilimanjaro in submontane to lower montane Croton-Caloden-
drum, Cassipourea and Ocotea forests, from 1340 to 2330 m.
Distribution: K 4; T 2, 3, 6; not known elsewhere. At Kilimanjaro
rare.
Pollination: insects, not further details available.
Luffa cylindrica (L.) M. Roem (Plate 11, 4e6).
Shape: spheroidal, circular ~100 mm.
Apertures: 3-colporate, long (5/6) colpi with large, elliptic broad
pori, colpi badly defined especially in other than equatorial view.
Exine: reticulate, heterobrochate, rather dense/small-meshed
reticulum.
Habitus: herbaceous climber or trailer to 15 m.
Habitat: persisting in old cultivations and near habitations,
frequently becoming naturalized in forest, woodland, bushland,
thicket and grassland; 0e1530 m. At Kilimanjaro in lower montane
gorge forest, roadsides, 900e1590 m.
Distribution: U 2e4; K 4, 7; T 1, 3, 4, 6, 8; widely distributed in
the Tropics and Subtropics as an escape from cultivation, doubtfully
a native of tropical East Africa. At Kilimanjaro scattered.
Pollination: insects, especially ants, but also other Hymenoptera,
as well as Coleoptera, Lepidoptera, and Diptera (Agarwal and
Rastogi, 2008).
Momordica boivinii Baill (Plate 11, 7e9).
Shape: subprolate ~61  72 mm, circular.
Apertures: 3-colporate, long (3/4) and narrow colpi, colpi
tapering towards poles and with thicker margo at the equator, pori
elliptic tall.
Exine: tectate, retipilate/reticulate, high and rather dense/small-
meshed reticulum, homobrochate?
Habitus: Herb climbing or trailing to 2 m from a perennial tu-
berous rootstock. Stems annual, herbaceous.
Habitat: Deciduous bushland, thicket, woodland and wooded
grassland; 0e1650 m. At Kilimanjaro in savanna woodland with
Combretum molle, Ozoroa insignis and Dombeya rotundifolia, from
900 to 1270 m.
346 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
Plate 11. Cucurbitaceae: 1e3 Diplocyclos schliebenii, 4e6 Luffa cylindrica, 7e9 Momordica boivinii, 10e11 Momordica foetida, 12 Oreosyce africana.
Distribution: U 1; K 1, 2, 4, 6, 7; T 2, 3, 5e8; Eastern Africa from
southern Ethiopia to South Africa (Natal). At Kilimanjaro very rare.
Pollination: by specialized oil bees of the genus Ctenoplectra
(Schaefer and Renner, 2010, 2011).
Momordica foetida Schumach (Plate 11, 10e11).
Shape: subprolate ~64  86 mm, circular.
Apertures: 3-colporate, colpi long and narrow with margo, pori
inconspicuously lalongate.
Exine: tectate, reticulate, very thick muri and small lumina and
reticulum flat compared to M. bovinii.
Habitus: perennial herb with woody rootstock. Stems trailing or
climbing up to 4.5 m, herbaceous, when old becoming rather
woody and rooting at the nodes.
Habitat: forest edges and clearings; margins of swamp and
riverine forest and of secondary thickets, also a weed and colonizer
of disturbed ground and of old cultivations; 0e2350 m. At Kili-
manjaro in riverine forests, forest clearings and edges, Chagga
homegardens, from 960 to 2180 m.
Distribution: U1e4; K1e6; T1e8; Z; widely distributed in
tropical Africa and in South Africa. At Kilimanjaro scattered.
Pollination: by specialized oil bees of the genus Ctenoplectra
(Schaefer and Renner, 2010, 2011).
Oreosyce africana Hook f (Plate 11, 12 & Plate 12, 1).
Shape: circular, spheroidal ~75e91 mm.
Apertures: 3-porate, large pori Ø ~15 mm with thin annulus and
small vestibulum.
Exine: tectate, psilate.
Habitus: climber or trailer to 3 m.
Habitat: upland grassland, also edges and clearings in rain-,
swamp and other groundwater forest and upland bamboo thicket;
900e3000 m. At Kilimanjaro in scattered in nearly all forest types
from the savanna to the upper montane zone, on forest clearings
and edges, on roadsides, fallow arable fields, waste places and
Chagga homegardens, from 900 to 3170 m.
Distribution: U 2e4; K 1e4; T 1e8; Bioko I., Cameroon Mt.,
Eastern DR Congo Republic, Ethiopia, Southern tropical Africa and
Madagascar. At Kilimanjaro widespread but in low frequencies.
Pollination: insects.
 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386 347

Plate 12. CUCURBITACEAE: 1 Oreosyce africana, CYPERACEAE: 2e3 Carex conferta, 4e6 Carex vallis-rosetto, 7e9 Cyperus amauropus, 10e11 Cyperus rigidifolius, DRACAENACEAE:
12e13 Dracaena afromontana, 14e15 Dracaena fragrans, EBENACEAE: 16e18 Euclea divinorum, ERICACEAE: 19e21 Agauria salicifolia, 22e24 Erica arborea, Euphorbiaceae: 25 Clutia
abyssinica var. abyssinica.

5.2.30. Cyperaceae or ± spheroidal. Most commen aperture type is one ulceroid

Small to tall perennial or annual herbs, rarely dwarf shrubs,
shrubs, or lianas, terrestrial or halophytic, rarely aquatic, epiphytic,
or epilithic. A cosmopolitan family with greatest diversity in humid
and semihumid tropics, but also often dominant in temperate and
cold temperate regions of the world, comprising 104 genera and
over 5000 species, of which 2000 belong to the genus Carex
(Goetghebeur, 1998).
5.2.30.1. Pollination. Cyperaceae are clearly adapted to anemophily.
Nevertheless, scattered species in various genera are known to be
visited by pollen-gathering insects or have traits indicating polli-
nation by insects (and other invertebrates). These include the
Cyperoideae species of Bolboschoenus, Cyperus, Eleocharis, and
Ficinia, and in the Caricoideae the genera Carex and Cymophyllus.
Many of these species are wind-pollinated as well, and even self-
pollination definitely occurs (Goetghebeur, 1998).
5.2.30.2. Pollen morphology. Grains are 1e4 aperturate, elongate
aperture at the thick end and three lateral, faintly marked poroid or
elongate apertures. Exine usually thin. Sexine as thick as nexine or
usually thicker (Erdtman, 1952).
Carex conferta Hochst. ex A. Rich (Plate 12, 2e3).
Shape: concial, longest axis ~53 mm, heteropolar.
Apertures: one ulceroid aperture at the thick end and three
indistinct lateral elliptic poroid regions.
Exine: thin, scabrate.
Habitus: perennial herb with tufts 0.15e0.45 m tall from a
mostly long-creeping rhizome, usually well spaced and ±slender.
Habitat: swamps, bogs, streamsides and moist ground in
bamboo, Hagenia, Hypericum and Erica zones, moorland and upland
forest, grassland with Acacia; 2250e3650 (-?3750) m. At Kili-
manjaro on moist ground in subalpine Erica bushland, from 2760 to
2870 m.
Distribution: U 2, 3; K 1, 3e5, T 2, 7; E DR Congo-Kinshasa,
Rwanda, Ethiopia. At Kilimanjaro rare.
Pollination: not known, probably insect and/or wind pollinated.
348 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
Carex vallis-rosetto K. Schum (Plate 12, 4e6).
Shape: ovoid ~65 mm, heteropolar.
Apertures: one ulceroid aperture at the thick end and four
indistinct lateral elongate poroid regions.
Exine: thin, thicker than in C. conferta, scabrate.
Habitus: tufted perennial with thick rhizome, 0.5e2 m tall.
Habitat: damp or swampy places in forest or forest edges of
Hagenia bamboo etc., Erica zone, riverine forest; 1000e3300 m. At
Kilimanjaro mainly in upper montane Hagenia riverine forest,
furthermore in Podocarpus, Juniperus and Erica excelsa forest, from
2350 to 3240 m.
Distribution: K 3e4; T 2e3, 6, 7; not known elsewhere. At
Kilimanjaro in the upper forest belt scattered.
Pollination: not known, probably insect and/or wind pollinated.
Cyperus amauropus Steud (Plate 12, 7e9).
Shape: heteropolar, spheroidal to ovoid ~35 mm, circular, often
folded.
Apertures: one ulceroid aperture at the thick end and four
lateral elongate poroid regions.
Exine: strongly scabrate to verrucate.
Habitus: perennial, fairly robust, succulent, up to 0.7 m tall.
Culms tufted, 0.15e0.6 m long.
Habitat: in grassland and wooded grassland, on rocky hills, and
on shallow soil covering rocks; 450e2100 m. At Kilimanjaro on
sun-exposed cliffs in the savanna and submontane cultivation zone,
growing together with Actiniopteris raddiata, A. semiflabellata,
Cyperus amauropus, Crassula volkensii and Xerophyta spekei,
furthermore in savanna grasslands, from 1010 to 1330 m.
Distribution U 1, 2; K 1e4, 6, 7; T 1e7; Rwanda, Sudan, Eritrea,
Ethiopia, Somalia, Zambia. At Kilimanjaro in the savanna scattered.
Pollination: not known, probably insect and/or wind pollinated.
Cyperus rigidifolius Steud (Plate 12, 10e11).
Shape: conical, longest axis ~35 mm, heteropolar.
Apertures: one ulceroid aperture at the thick end and 5e6
distinct lateral mostly very long conical colpoid regions, aperture
shape very variable in this species.
Exine: thin, stratification not visible, psilate.
Habitus: perennial, slender to fairly robust, with a woody base
and curving horizontal stolons up to 15 cm long, up to 72 cm tall.
Habitat: in seasonally wet grassland, swamps, bushland;
1700e2800 m. At Kilimanjaro in wet submontane and lower
montane meadows, in swamps on streamsides, from 1250 to
1880 m.
Distribution: U 2e4; K 1, 3e6; T 1e4, 7; DR Congo-Kinshasa,
Rwanda, Burundi, Ethiopia, South Africa, Swaziland. At Kili-
manjaro in the cultivation belt below the forest scattered.
Pollination: not known, probably insect and/or wind pollinated.
5.2.31. Dracaenaceae
Trees, shrubs, sometimes scandent, unbranched, suffrutices or
rhizomatous geophytes, sometimes succulent, glabrous, from less
than 10 cm to over 40 m tall. A family of one genus of some 100
species native to the warmer regions of the Old World, odd species
in Hawaii, Central America, Cuba and Macaronesia. Widely intro-
duced and established in tropical climates elsewhere (Bos, 1998).
5.2.31.1. Pollination. Nocturnal flowering, strong fragrance during
the late evening, and copious production of nectar on the inflo-
rescence point to pollination by night-active animals. In Dracaena
spontaneous self-pollination is quite possible and has successfully
been effected artificially on several occasions (Bos, 1998).
5.2.31.2. Pollen morphology. Pollen grains are sulcate, either
microechinate or not. Tectate, sexine thicker than nexine (Erdtman,
1952); fossulate, reticulate (Dracaena) or psilate (Sanseveria).
Dracaena afromontana Mildbr (Plate 12, 12e13).
Shape: sub-prolate ~44  69 mm to spheroidal, ± circular
~57 mm.
Apertures: monocolpate, colpus length 5/5, wide with smooth
margins.
Exine: tectate, finely reticulate, heterobrochate (Note: Sculp-
turing maybe difficult to identify due to poor preservation).
Habitus: shrub or tree, 2e12 m high, sometimes straggling. Stem
to 25 cm in diameter, branches few.
Habitat: rain-forest, moist forest, or bamboo forest, often
forming dense stands in the understorey; 1600e2700 m. At Kili-
manjaro in the whole sub to middle montane forest belt, rarely in
upper montane Podocarpus and Juniperus forest; 1280e2780 m.
Distribution: U 1e3; K 1, 3e4, 6e7; T 2e4, 6e7; DR Congo,
Rwanda, Burundi, Ethiopia, Malawi. At Kilimanjaro widespread in
the forest belt.
Pollination: probably night active animals or self-pollination
(Bos, 1998).
Dracaena fragrans (L.) Ker Gawl (Plate 12, 14e15).
Shape: (sub)prolate ~44  66 mm, ± circular.
Apertures: monocolpate, colpus length 5/5, wide (more narrow
than in D. afromontana) with smooth margins.
Exine: tectate, reticulate, reticu-pilate (Note: Sculpturing maybe
difficult to identify due to poor preservation).
Habitus: shrub or tree, 1.5e15 m high. Stem single, up to 30 cm
in diameter, or in forest often with a mass of horizontal stems near
the ground from which vertical stems arise.
Habitat: moist forest, often near streams; may be locally com-
mon and may form thickets; 600e2250 m. At Kilimanjaro in colline
to lower montane riverine and gorge forests, widely planted in the
Chagga Homegardens; 980e1770 m.
Distribution: U 2e4; K 1e7; T 1e8; throughout tropical Africa. At
Kilimanjaro, frequent in the submontane and lower montane forest
and Homegardens.
Pollination: from field observations it is assumed that D. fragans
is pollinated by moths (Bergsdorf, 2006).
5.2.32. Ebenaceae
Mostly evergreen dioecious, rarely monoecious or polygamous
trees or shrubs. A family comprising two genera and ca. 500e600
species, distributed mainly in the tropical and subtropical regions
of both the Old and the New World (Wallno € fer, 2004).
5.2.32.1. Pollination. The flowers of various species are reported to
be visited by bees and other insects (Wallno€ fer, 2001, 2004).
5.2.32.2. Pollen morphology. Pollen grains are 3-colporate and
prolate-spheroidal to prolate. The sexine is as thick as the nexine or
thicker. Grains are psilate or finely scabrate. Pori are mostly well
defined, usually lalongate, sometimes with indistinct lateral edges
or rarely indistinguishable. The pollen is remarkably constant
throughout the family, and the main variations are in size and
shape of the grains and the apertures (Erdtman, 1952; Wallno €fer,
2004; Geeraerts et al., 2009).
Euclea divinorum Hiern (Plate 12, 16e18).
Shape: radially symmetric, subprolate to prolate ~22  30 mm,
trilobate.
Apertures: 3-colporate, long (4/5) slightly open colpi with
smooth margins, circular (sometimes oval) pori with irregular
margins.
Exine: tectate, exine thick, psilate, granulate, perforate.
Habitus: evergreen shrub or small tree 1.8e9 (15) m tall.
Habitat: grassland, mostly with scattered trees, open bushland,
thicket on anthills, secondary forest, margins of evergreen forest,
often on stony slopes; 0e2700 m. At Kilimanjaro in drier forest
 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
communities, from savanna riverine forests, submontane Croton-
Calodendrum forest to lower montane Cassipourea forests;
1080e2030 m.
Distribution: U 1e4; K 1, 3e7; T 1e7; widely distributed from
Ethiopia to South Africa and Namibia, also in Socotra. At Kili-
manjaro scattered.
€ fer, 2001).
Pollination: probably various insects (Wallno
5.2.33. Ericaceae
Evergreen or deciduous shrubs, rarely scandent, lianas, or trees.
A mainly temperate, warm temperate and montane tropical family
of some 124 genera and 4100 species (Stevens et al., 2004).
5.2.33.1. Pollination. Self-pollination and cross-pollination
wind, insects and birds (Sarwar, 2007, and literature herein).
5.2.33.2. Pollen morphology. Grains are usually in radiosymmetric,
triangular to globular tetrahedral tetrads 23e82 mm in diameter.
They are usually 3-colporate, sometimes 3-colpate or tricolpor-
idate, and with definite transverse furrows. Aperture number can
vary from 3 to 5 at the species level in Enkianthus. The exine is
scabrate, verrucate, microrugulate or psilate (the latter especially
by aperture margins) (Erdtman, 1952; Stevens et al., 2004; Sarwar,
2007). The pollen morphology of the genus Erica L. and some rel-
atives has been studied in detail by Sarwar and Takahashi (2014).
Agauria salicifolia (Comm. ex Lam.) Hook. f. ex Oliv (Plate 12,
19e21). SYN Agarista salicifolia (Comm. ex Lam.) G. Don.
Shape: tetrad Ø ~27 mm, isodynamosporus (all grains of the
same size), single grains lobate ~20 mm.
Apertures: 3-colporate, evenly narrow colpi with distinct costae,
colpus membrane granulate, narrow lalongate pores.
Exine: tecate, invisible collumella, mesocolpial exine thicker
than apocolpial exine, psilate or finely granulate.
Habitus: evergreen shrub or tree 0.9e20 m, much branched.
Bole occasionally large, up to 0.6 m across.
Habitat: dry and moist forest, secondary forest, forest margins,
evergreen bushland to woodland or clumped or scattered tree
grassland, often on rocky slopes, heath zone; fire-resistant; may be
locally common or dominant, belt-forming in Virunga; (1050-)
1550e3500 m. At Kilimanjaro bimodal distribution, avoiding the
wettest parts of the middle montane Ocotea forests, occurring in
lower montane Ocotea forests and upper montane Podocarpus, Erica
and Juniperus forests, also as a remnant tree in (formerly burnt)
subalpine ericaceous bushland; 1670e3000 m.
Distribution: U 1e3; K 1e7; T 2e8; widespread in tropical Africa
from Cameroon to north-east Africa and south to Zambia,
Madagascar (Mascarene I.). At Kilimanjaro frequent.
Pollination: probably wind, birds and insects (Rebelo and
Siegfried, 1985).
Erica arborea L (Plate 12, 22e24).
Shape: oblate tetrad Ø 30 mm, isodynamosporus (all four grains
of the same size, Erdtman, 1952), single grains triangular/trilobate
~19 mm.
Apertures: 3-colporate, narrow colpi with distinct costae, colpi
often wider in the pore area, lalongate pori.
Exine: tectate, invisible collumella, mesocolpial exine thicker
than apocolpial exine, rugulate to rugulate-granulate.
Habitus: shrub or tree 0.3e7.5 m high.
Habitat: dominant in a zone above the forest on many moun-
tains, and probably a pyrophyte; also in open forest, bamboo,
grassland, and on upper moorland, often in rocky places; often
together with Stoebe, Artemisia, Faurea or Protea; (1600-)
2000e3900(-4500) m. At Kilimanjaro in subalpine ericaceous
bushland; 2640e4000 m.
Distribution: U 1e3; K 1e6; T 2, 3; Chad (Tibesti), Kongo
349
(Kinshasa), Sudan, Ethiopia, Somalia, North Africa and Europe from
the Canaries and N. Spain to the Black Sea region and Saudi Arabia
and Yemen. At Kilimanjaro scattered and rarer than the other two
occurring Erica trees (E. excelsa and E. trimera). Palynologically, the
three Erica species are not (easily) distinguishable.
Pollination: probably self-pollination and cross-pollination by
wind, insects and birds (Sarwar, 2007, and literature herein).
5.2.34. Euphorbiaceae
Monoecious or dioecious trees, shrubs, or herbs, sometimes
succulent or scandent. Webster (2014) contrues to include 299
genera with 8000 species. Although the family is nominally
cosmopolitan, it is poorly represented in cool temperate zones and
by best developed in subtropical and tropical regions.
5.2.34.1. Pollination. Euphorbiaceae show a great diversity of
pollination systems and pollinator species. Pollination occurs
abiotically via wind in Acalypha, some Macaranga and Mallotus
species, Mercurialis, Ricinus and probably many other Acalypheae.
Wind pollination has also been reported in Bernardia Bertya,
“Celaenodendron” some Croton species and possibly in two or more
species of Phyllanthus. Species with biotic pollination are visited by
a variety of pollinator species seeking nectar, particularly those
belonging to the insect orders Hymenoptera and Diptera. These
plants include Croton, Jatropha and Tragia. Specialized pollination is
seen in some species of Macaranga, Breynia, Glochidion, and some
Phyllanthus (Webster, 2014, and literature herein).
5.2.34.2. Pollen morphology. Pollen features have been used as
diagnostic characters to distinguish the seven subfamilies: Phyl-
lanthoideae (tribe Wielandieae, Phyllantheae): prolate 3-colporate
grains with reticulate (semitectate) exine, perforate or discontin-
uous ectexine with distinct columellae and a continuous homoge-
neous endexine. Oldfieldioideae: grains mostly oblate-globose,
apertures contracted, from brevicolpate to porate and from zono-
porate to pantoporate, four apertures to pantoporate, exine is
modified to radially elongate, tapering spines. Peroideae: palyno-
logically diversified. Cheilosoideae: characterized by echinate
tectum. Acalyphoideae: 3-colporate but with great diversity in
structure and sculpture. Crotonoideae: mostly inaperturate with an
exine sculpture of triangular (rounded) structures in continuous
arraysd now widely known as “Croton pattern”; also colporate,
colpate and porate. Euphorbioideae: 3-colporate; tribe Euphor-
biaeae: conspicuous intine bands bordering the colpi (Webster,
2014, and literature herein).
Clutia abyssinica Jaub. & Spach var. abyssinica (Plate 12, 25 &
Plate 13, 1e2).
Shape: circular ~44 mm, (sub)prolate ~44  60 mm.
Aperture: 3-colporate, syncolpate, colpi with margo, tall elliptic
pori with costa.
Exine: collumellate, tectate, reticulate.
Habitus: large erect woody herb, shrub or small tree up to 8 m
tall with straight branches.
Habitat: widespread and common at forest edges, associated
bushland and wooded grassland, secondary associations (often an
indicator of old forest areas), lakesides, riverine, evergreen thickets;
(1000-)2000e3700 m. At Kilimanjaro showing a bimodal distri-
bution pattern, occurring in heavily degraded lower montane
bushlands and upper montane Juniperus forests; 1170e2990 m.
Distribution: U 1e4; K 1e7; T 1e7; Sudan to Somalia and south
to Zaire, Angola and South Africa (Natal). At Kilimanjaro scattered.
Pollination: not known.
Croton megalocarpus Hutch (Plate 13, 3e4).
Shape: circular, spheroidal ~60 mm.
Aperture: inaperturate.
350 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386

Plate 13. EUPHORBIACEAE: 1e2 Clutia abyssinica var. abyssinica, 3e4 Croton megalocarpus, 5 Euphorbia engleri, 6e8 Euphorbia schimperiana var. schimperiana, 9e10 Macaranga
kilimandscharica, 11e12 Phyllanthus boehmii var. boehmii, FABACEAE: 13e15 Aschynomene schimperi, 16e17 Calpurnia aurea, 18e20 Crotalaria axillaris, 21e23 Crotalaria goodiformis,
24e26 Crotalaria keniensis, 27e28 Dalbergia lactea, 29e31 Desmodium repandum, 32e34 Eriosema montanum var. montanum, 35e37 Erythrina burtii.

Exine: reticulate, ‘crotonoid’. forests, mainly in lower to middle montane Cassipourea and Ocotea
Habitus: a large monoecious or occasionally dioecious tree up to forests and upper montane Erica and Juniperus forests;
35 m tall, though commonly 15e25 m. 960e2780 m.
Habitat: evergreen forest; (700-)1200e2400 m. At Kilimanjaro Distribution: U 3; K 1, 3e7; T 2; not known elsewhere. At Kili-
in colline riverine forests, submontane Croton-Calodendrum forests manjaro frequent.
and lower montane Cassipourea forests; 960e1980 m. Pollination: not known; probably insects.
Distribution: U 2, 4; K 1, 3e7; T 1e4, 6, 8; Zaire (Orientale, Kivu Euphorbia schimperiana Scheele var schimperiana (Plate 13,
and Shaba), Rwanda, Burundi, Zambia, Malawi, Mozambique. At 6e8).
Kilimanjaro frequent. Shape: subprolate to spheroidal ~40  45 mm, circular to
Pollination: wind and/or insects. trilobate.
Euphorbia engleri Pax (Plate 13, 5). Apertures: 3-colporate, colpi 4/5 with acute ends, colpus mar-
Shape: spheroidal to subprolate ~33  36 mm, circular. gins narrow, slightly thickened at the pori, colpus membrane
Apertures: 3-colporate, lalongate pori, colpi (4/5) relatively granular, pori lalongate.
narrow with smooth margins, thickened at the pori, colpus mem- Exine: tectate, microreticulate becoming reticulate along the
brane granular. colpal margin especially towards the pori, heterobrochate.
Exine: tectate, indistinctly perforate, reticulate, homobrochate. Habitus: annual or short-lived perennial herb, erect to 2 m.
Habitus: shrubby, perennial herb to 3 m. Habitat: in grasslands, forest edges and clearings and a weed of
Habitat: forest undergrowth and dense bushland; land cleared for cultivation; 1100e3350 m. At Kilimanjaro mainly in
1600e2700 m. At Kilimanjaro rarely in colline to submontane upper montane Erica, Hagenia and Juniperus forests and clearings;
 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
1930e3090 m.
Distribution: U 1e4; K 1e6; T 2e4, 6, 7; the Arabian Peninsula,
N. Somalia and Ethiopia; westwards to Zaire and Cameroon; and
south throughout Malawi and into Zimbabwe. At Kilimanjaro
scattered.
Pollination: not known; probably insects.
Macaranga kilimandscharica Pax (Plate 13, 9e10).
Shape: subprolate ~20  24 mm, spheroidal to trilobate.
Aperture: 3-colporate, lalongate narrow pori, colpi (4/5) narrow
with thin costa.
Exine: tectate, tectum thick and perforate with microspinules,
columellate, faintly scabrate.
Habitus: small to medium tree 4.5e18 m, or large tree up to
27 m.
Habitat: evergeen forest, locally common to dominant in wetter
types, regenerating profusely in clear-felled areas, in secondary
forest, on forest edges, in riverine forest and disturbed places;
1310e3000 m. At Kilimanjaro from lower to middle montane
riverine, Cassipourea and Ocotea forests; 1640e2350 m.
Distribution: U 1e3; K 2e5; T 2e4, 6, 7; Sudan, Ethiopia, Zaire,
Rwanda, Burundi, Zambia, Malawi; At Kilimanjaro widespread.
Pollination: wind, or maybe thrips (Thusanoptera) (Moog et al.,
2002).
Phyllanthus boehmii Pax var. boehmii (Plate 13, 11e12).
Shape: prolate ~15  31 mm, circular.
Aperture: 3-colporate, lalongate pori with costa to nearly
zonoporate, narrow colpi (4/5).
Exine: tectate, reticulate, homobrochate.
Habitus: erect annual herb up to 0.4 m tall, sometimes getting
woody at the base and then reaching 2 m in height, or else a pro-
cumbent perennial.
Habitat: damp places in forest, grassland and woodland;
1050e2450 m. At Kilimanjaro mainly in submontane to lower
montane grasslands and forest tracks; 1680e2310 m.
Distribution: U 1, 2; K 3, 4, 6; T 1, 4, 6, 7; Ethiopia, Malawi,
Zambia. At Kilimanjaro widespread.
Pollination: probably moths (Kawakita and Kato, 2004), maybe
wind and other insects (Webster, 2014).
5.2.35. Fabaceae
Trees, or shrubs, or herbs, or lianas. About 430 genera and 9000
species with a cosmopolitan distribution (Watson and Dallwitz,
1992 onwards).
5.2.35.1. Pollination. Entomophilous, or ornithophilous, or cheir-
opterophilous (Watson and Dallwitz, 1992 onwards).
5.2.35.2. Pollen morphology. Usually 3-colporate (sometimes pro-
vided with 2,4, or 6 apertures), ranging in shape from perprolate to
prolate (Erdtman, 1957).
Aeschynomene schimperi Hochst. ex A. Rich (Plate 13, 13e15).
Shape: prolate ~19  29 mm, subtriangular.
Apertures: 3-colporate, colpi long and (5/6) wide at the equator
with acute ends at poles, pori lolongate/elliptic tall without distinct
margins.
Exine: tectate, reticulate, homobrochate.
Habit: shrubby aromatic herb or small shrub, 0.6e3 m tall,
usually erect but rarely straggling.
Habitat: swampy places, stream banks, lake sides and rock pools
in grassland or wooded areas, often actually standing in water;
60e2340 m. At Kilimanjaro in colline to submontane swamps on
lake shores and streamsides; 1070e1500 m.
Distribution: U 1e4; K 1, 3, 4, 6, 7; T 1e5, 7, 8; widespread
throughout tropical Africa from Senegal and Ethiopia to Zimbabwe,
also in Madagascar. At Kilimanjaro rare.
351
Pollination: in glasshouse conditions A. schimperi has been
observed to be selfpollinating (Loch and Ferguson, 1999). There are
no published observations from the field. Bees have been observed
to pollinate other Aeschynomene species (Carleial et al., 2015).
Calpurnia aurea Baker (Plate 13, 16e17).
Shape: subprolate ~24  31 mm, circular to slightly trilobate.
Apertures: 3-colporate; colpi (5/6) wide, colpus margins
smooth, furrows maybe smooth or crustate, pori inconspicuous.
Exine: tectate, tectum perforate, reticulate, homobrochate,
exine thickness between 0,5 and 1 mm.
Habitus: bush or small tree 2e10 m tall.
Habitat: margins and clearings of upland rain-forest and
riverine forest; 1300e2250 m. At Kilimanjaro in submontane
Croton-Calodendrum forests and clearings; 1470e1480 m.
Distribution: U 1, 3; K 3e5; T 2e4; highlands of eastern tropical
Africa north to Eritrea and as far west as the Central African Re-
public, eastern DR Congo and western Angola, extending south to
the eastern Cape Province of South Africa. At Kilimanjaro rare.
Pollination: not known; bees have been observed on the
flowers. Bees are also pollinators of other Calpurnia species
(Leistner, 2000).
Crotalaria axillaris Aiton (Plate 13, 18e20).
Shape: prolate to slightly rectangular ~17  29 mm, trilobate to
circular.
Apertures: 3-colporate, colpi long (5/6), narrow, slightly con-
stricted at equator with acute ends, large pori rectangular or elliptic
broad.
Exine: tectate, micro-reticulation fine and faint, homobrochate.
Habitus: shrub or woody herb up to 1e4 m tall.
Habitat: forest margins, deciduous woodland and bushland,
grassland, also in abandoned cultivations, secondary scrub and on
coral crags; 0e2250 m. At Kilimanjaro in colline to submontane
riverine and Croton-Calodendrum forests; 980e1850 m.
Distribution: U 2; K 3e7; T 2e4, 6; Z; Ghana, SW. Ethiopia, DR
Congo, Mozambique, Malawi, Zambia, Angola. At Kilimanjaro
scattered.
Pollination: not known for this species. Other Crotalaria species
have been observed to be pollinated by bees and can be self-
pollinating (Etcheverry, 2001; Jacobi et al., 2005).
Crotalaria goodiformis Vatke (Plate 13, 21e23).
Shape: subprolate to rectangular ~19  29 mm, spheroidal to
slightly trilobate.
Apertures: 3-colporate, colpi long (5/6) and narrow, pori rect-
angular to elliptic broad.
Exine: tectate, micro-reticulation very fine and faint,
homobrochate.
Habitus: bushy shrub or subshrub, sometimes scandent, up to
2.6 (?-4.5) m tall.
Habitat: margins and clearings of lowland and upland rain-
forest, dry evergreen forest, deciduous woodland and bushland,
wooded grassland, also persisting on abandoned cultivations;
75e2100 m. At Kilimanjaro in colline to submontane riverine and
Croton-Calodendrum forests; 900e2090 m.
Distribution: K 1, 4, 6, 7; T 1e8; Mozambique, Eastern DR Congo.
At Kilimanjaro scattered.
Pollination: not known for this species. Other Crotalaria species
have been observed to be pollinated by bees and can be self-
pollinating (Etcheverry, 2001; Jacobi et al., 2005).
Crotalaria keniensis Baker f (Plate 13, 24e26).
Shape: subprolate ~16  19 mm, slightly trilobate to circular
~19 mm.
Apertures: 3-colporate, colpi long (4/5) and narrow, pori rect-
angular to elliptic broad.
Exine: tectate, micro-reticulation very fine and faint.
Habitus: bushy or weakly straggling woody herb or shrub, rarely
352 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
climbing, up to 2 (4) m tall.
Habitat: margins and clearings of upland rain-forest, upland
evergreen bushland and grassland, very common in elevated vol-
canic soils in the region of the Great Rift Valley; 1500e2700 m. At
Kilimanjaro in clearings in submontane Croton-Calodendrum for-
ests and upper montane Juniperus forests; 1890e2450 m.
Distribution: U 3; K 1, 3e6; T 2, 7; SW Ethiopia. At Kilimanjaro
scattered.
Pollination: not known for this species. Other Crotalaria species
have been observed to be pollinated by bees and can be self-
pollinating (Etcheverry, 2001; Jacobi et al., 2005).
Dalbergia lactea Vatke (Plate 13, 27e28).
Shape: subprolate ~21  25 mm, circular to slightly trilobate
~21 mm.
Apertures: 3-colporate, colpi long (4/5) and slightly open with
smooth margins, constricted at equator, elliptic to rectangular
broad (faint) pori.
Exine: tectate, microreticulate, homobrochate, sexine thicker
than nexine.
Habitus: small tree or shrub, generally scandent if support
available, 3e9 (25) m tall, ± evergreen.
Habitat: margins and clearings of rain- and swamp-forest,
riverine forest and bushland, upland evergreen bushland and
grassland; 540e2400 m. At Kilimanjaro mainly in colline to sub-
montane riverine forests and lower montane Cassipourea and
Ocotea forests; 990e2130 m.
Distribution: U 1e4; K 2e5, 7; T 1e4, 6e8; widespread in
tropical Africa from Nigeria to Ethiopia and south to Mozambique
and Zimbabwe. At Kilimanjaro widespread.
Pollination: not known for this species. However, D. sissoo is
known to be cross- and self-pollinated by honey bees, beetles,
butterflies and thrips (Vasudeva and Sareen, 2009).
Desmodium repandum (Vahl) DC (Plate 13, 29e31).
Shape: spheroidal ~20 mm, convex triangular.
Apertures: 3-colporate, broad, badly defined, rectangular pori,
open and long colpi (4/5), slightly recessed with granular margins.
Exine: collumellate, tectate, perforate, foveolate.
Habitus: perennial herb or slender shrub with a woody base,
climbing, scrambling, or forming dense undergrowth, 0.5e1 m
high.
Habitat: occasional in shady grassland and marginal areas of
cultivated land, but more often one of the dominant components of
shaded places in dry evergreen forest or rain-forest or at the forest
margin; 1000e3000 m. At Kilimanjaro in colline to submontane
riverine forests, submontane Croton-Calodendrum forests, lower to
middle montane Cassipourea and Juniperus forests and lower
montane Ocotea forests, also in Chagga homegardens and open
habitats (roadsides, swamps); 970e2450 m.
Distribution: U 1e4; K 1e7; T 1e8; tropical and subtropical
Africa, Madagascar, Mascarene Is., India, Malesia. At Kilimanjaro
widespread.
Pollination: various bee species (Gikungu, 2006). Probably also
self-pollinating like other Desmodium species (Alem an et al., 2014).
Eriosema montanum var. montanum Baker f (Plate 13, 32e34).
Shape: subprolate ~32  39 mm, circular to trilobate.
Apertures: 3-colporate, long (4/5 to 5/6) and open colpi recessed
with acute ends, slightly constricted at equator large circular pori.
Exine: tectate, reticulate, heterobrochate, reticulation absent or
faint along colpal margins, sexine thicker than nexine.
Habitus: perennial herb, often subshrubby, 0.2e1.5 m tall.
Habitat: grassland, scrub, bushland, forest edges; 900e2520 m.
At Kilimanjaro in dry submontane and lower montane grasslands
and forest edges; 1310e1790 m.
Distribution: U 1e4; K 3e5, 7; T 1e4, 6e8; Nigeria, Cameroon,
Ethiopia, DR Congo, Malawi, Zambia, Zimbabwe, Angola. At
Kilimanjaro scattered.
Pollination: not known for this species. Probably self-pollinating
and/or pollination by bees, flies, butterflies and wasps like other
Eriosema species (SGRP; Johnson et al., 2009).
Erythrina burtii Baker f (Plate 13, 35e37).
Shape: spheroidal~46 mm, (sub)triangular.
Apertures: 3-colporate, colpi (3/5) open with acute ends, pori
circular.
Exine: tectate, sexine thicker than nexine, reticulate with
medium-sized lumina, heterobrochate, lumina size smaller around
apertures.
Habitus: flat-topped tree 3.5e15 (18) m tall.
Habitat: Acacia, Combretum or Acacia, Commiphora bushland or
derived grassland; 800e1650 m.
Distribution: K 1, 3, 4, 6, 7; T 1e3, 5, 6; not known elsewhere.
Pollination: nektar feeding birds (Baker et al., 1998).
Glycine wightii (Graham ex Wight & Arn.) Verdc. agg (Plate 14,
1e3).
Shape: triangular (convex), subprolate to rhombic tall
~35  40 mm.
Apertures: 3-porate, angulaperturate, pori circular and large Ø
~8 mm.
Exine: tectate, sexine thicker than nexine, reticulate with large-
sized lumina and very broad muri, heterobrochate, lumina size
smaller around pori, sometimes pore area protruding.
Habitus: perennial climber or trailer, often woody at base,
0.6e4.5 m long.
Habitat: forest, grassland, grassland with scattered trees and
secondary grassland, scrub, thicket and woodland, sometimes in
cultivations; 0e29,000 m. At Kilimanjaro in dry savanna forest, in
colline and submontane riverine forests, Chagga homegardens,
open vegetation on roadsides, dry savanna and submontane
Hyparrhenia grasslands; 780e2000 m.
Distribution: U 1e4; K 1e6; T 1e8; Z; P; Arabia, Ethiopia, DR
Congo to S. and W Africa, Angola, Malawi, Zambia. At Kilimanjaro
widespread.
Pollination: honey bees (Nogueira-Couto et al., 1998).
Indigofera atriceps subsp. kaessneri (Bak. F.) Gillett (Plate 14,
4e6).
Shape: triangular/trilobate, spheroidal, ~27 mm.
Apertures: 3-colporate, recessed colpi (3/5) open with acute
ends and constricted at equator, pori inconspicuous.
Exine: tectate, perforate, psilate.
Habitus: erect herb up to 2 m tall.
Habitat: Grassland, bushland and forest margins; 900e2400 m.
At Kilimanjaro in dry colline savanna grasslands, submontane and
lower montane Hyparrhenia and Bulbostylis grasslands and subal-
pine Festuca obturbans tussock grasslands; 1150e2640 m.
Distribution: U 1e3; K1e5, 7; T 1e4; S Ethiopia, NE Zambia. At
Kilimanjaro widespread.
Pollination: not known for this species. Bees generally polli-
nated Indigofera sp. (Martins, 2014).
Indigofera swaziensis Bolus subsp. swaziensis (Plate 14, 7e8).
Shape: spheroidal ~28 mm, triangular.
Apertures: 3-colporate, colpi (4/5) open with margo and acute
ends, circular to elliptic tall pori.
Exine: tectate, reticulate, heterobrochate, no or faint reticulation
around aptertures.
Habitus: softly woody shrub up to 3 m tall.
Habitat: upland evergreen forest margins and bushland;
1200e2450 m. At Kilimanjaro mainly in dry colline savanna forests,
submontane Croton-Calodendrum forests and montane Juniperus
forests; 1190e2400 m.
Distribution: U 1; K 1e6; T 1, 2; southern Sudan, South Africa,
Swaziland. At Kilimanjaro scattered.
 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386 353

Plate 14. FABACEAE: 1e3 Glycine wightii agg., 4e6 Indigofera atriceps subsp. kaessneri, 7e8 Indigofera swaziensis subsp. swaziensis, Flacourtiaceae: 9e12 Dasylepis integra, 13e15
Dovyalis abyssinica, 16e18 Rawsonia lucida, HAMAMELIDACEAE: 19e20 Trichocladus ellipticus, HYPERICACEAE (partly Clusiaceae): 21e23 Garcinia volkensii, 24e26 Hypericum
revolutum ssp. keniense, Icacinaceae: 27e28 Apodytes dimidiata, IRIDACEAE: 29e30 Gladiolus watsonioides, LAMIACEAE: 31e33 Clerodendron johnstonii, 34e36 Leonotis mollissima,
37e40 Leucas volkensii, 41e42 Plectranthus alboviolaceus, 43e44 Plectranthus laxiflorus.

Pollination: not known for this species. Bees generally pollinate
Indigofera sp. (Martins, 2014).
5.2.36. Flacourtiaceae
Trees or shrubs. Pantropical family represented by 21 genera
and 43 species in East Africa, out of a world total of about 85 genera
and approximately 1100 species (Watson and Dallwitz, 1992
onwards).
5.2.36.1. Pollination. Entomophilous (Watson and Dallwitz, 1992
onwards).
5.2.36.2. Pollen morphology. Pollen grains usually 3-colporate
(-colporidate), subprolate to prolate. Sexine usually as thick as
nexine or slightly thicker or thinner, often finely reticulate. Brochi
often decrease in size towards colpi. Pori lalongate (Erdtman, 1952).
Dasylepis integra Warb (Plate 14, 9e12).
Shape: subprolate, trilobate, ~20  22 mm.
Apertures: 3-colporate, colpi (4/5) open with acute ends,
constricted at equator, pori circular.
Exine: tectate, psilate.
Habitus: shrub or tree, upt to 12 (35) m tall. Trunk slender.
Habitat: understorey of rain-forest; 900e2200 m. At Kili-
manjaro mainly in submontane riverine and Croton-Calodendrum
forests and lower montane Cassipourea and gorge forests;
1280e2330 m.
Distribution: K 7; T 2, 3, 6; not known elsewhere. At Kilimanjaro
scattered.
Pollination: insects; no further details known.
Dovyalis abyssinica (A. Rich.) Warb (Plate 14, 13e15).
Shape: (sub)prolate ~20  28 mm, trilobate.
Apertures: 3-colporate, colpi (4/5) open with acute ends, pori
elliptic tall.
Exine: tectate, finely reticulate, homobrochate.
Habitus: shrub or tree, much branched, up to 8 m tall.
Habitat: upland rain-forest to riparian and dry evergreen forest,
sometimes in open wooded grassland; 1500e3000 m. At Kili-
manjaro in nearly all main forest types, in colline and submontane
354 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
riverine forests, Croton-Calodendrum forests, lower to middle
montane Cassipourea and Ocotea forests, upper montane Podo-
carpus and Juniperus forests as well as Chagga homegardens;
1010e2920 m.
Distribution: U 1, 3,4; K 1e7; T 2, 3, 5e7; Ethiopia, Somali Re-
public, Socotra, Malawi. At Kilimanjaro widespread.
Pollination: by stingless bees (Vit et al., 2013, p. 323), other bees
and flies (Martins, 2014).
Rawsonia lucida Harv. & Sond (Plate 14, 16e18).
Shape: spheroidal ~19 mm, circular to slightly trilobate.
Apertures: 3 colporate, colpi (4/5) wide with smooth margins
and acute ends, pori inconspicuous.
Exine: tectate, faintly and finely reticulate.
Habitus: shrub or tree, rarely up to 20 m tall.
Habitat: understorey and shrub layer of lowland and upland
rain-forest, dry evergreen forest, semi-swamp and riverine forest;
50e1900 m. At Kilimanjaro only in lower forest types, in colline and
submontane riverine forests and submontane Croton-Calodendrum
forests; 900e1500 m.
Distribution: U 2, 3; K 4e7; T 1e3, 6e8; P; Somalia, Sudan,
Angola, Zaire, Malawi, Zambia, Zimbabwe, Mozambique,
Swaziland, South Africa. At Kilimanjaro scattered.
Pollination: not known.
5.2.37. Hamamelidaceae
Trees or shrubs. The family has a wide but very scattered and
relictic distribution with the greatest diversity in Eastern Asia. It
contains four subfamilies, 30 genera with ca. 100 spp., 12 of them
monotypic (Endress, 1993).
5.2.37.1. Pollination. In most genera pollination has not been
studied in the natural environment. Insect and wind pollination
occurs (Endress, 1993, and literature herein).
5.2.37.2. Pollen morphology. Pollen grains in the main part of the
family are 3-colpate, 3-colporidate (occasionally 4-rupate or 6-
rugate), suboblate - subprolate. Average length of the longest axis
varies from 17 mm to 37 mm. Sexine about as thick as nexine or
slightly thicker or thinner. OL-pattern. Grains are tectate-
columellate, reticulate. Apertures covered by entirely or partially
granular membranes; the latter are sometimes ± operculoid. Pollen
grains in Liquidambaroideae are polyfoveolate. The apertures are
usually not of the same size and shape, nor are they spaced regu-
larly (Erdtman, 1952; Endress, 1993).
Trichocladus ellipticus Eckl. & Zeyh (Plate 14, 19e20).
Shape: spheroidal to suboblate ~12 mm, triangular/trilobate/
circular.
Apertures: 3-colp(orid)ate, colpi (4/5) open with smooth mar-
gins and obtuse ends, pori absent or inconspicous.
Exine: tectate, reticulate with relatively high reticulum,
homobrochate.
Habitus: evergreen shrub or tree, sometimes somewhat
scrambling, (2-) 5e10 (18) m tall.
Habitat: upland rain-forest, often dominant by streams and in
swampy places, also extending into riverine forest, Acacia lahai
woodland and swamp forest; 1140e2700 m. At Kilimanjaro in
colline and submontane riverine forests and submontane Croton-
Calodendrum forests; 1170e2090 m.
Distribution: U 1e4; K 1e6; T 1e3, 6, 7; DR Congo, Ethiopia,
Malawi, Mozambique, Zambia, Angola. At Kilimanjaro rare.
Pollination: concluding from flower structure, the genus Tri-
chocladus may attract especially small dipterans acting as pollina-
tors (Endress, 1989), however according to Kubitzki (1977)
Trichocladus is not adapted to fly pollination. Observations on
pollination are missing.
5.2.38. Hypericaceae ¼ Clusiaceae-Guttiferae
Evergreen shrubs or trees, epiphytic or not. A family of 27 genera
and about 1090 species; largely restricted to the lowland tropics
(Stevens, 2007).
5.2.38.1. Pollination. Bees, birds, butterflies, monkeys, bat and also
lemurs have reported to be pollinating agents; species are usually
specialized for one pollinator type (Stevens, 2007, and literature
herein).
5.2.38.2. Pollen morphology. Pollen grains are usually monads, 3-
colporate, sometimes triporate; Garcinieae and Symphonieae
often have more than three apertures. Costa colpi often present,
and there is considerable variation in endoapertue type and
orientation. Pollen surface maybe reticulate, rugulose, fossulate,
foveolate, psilate or scabrate; supratectal elements occur in some
general, notably some species of Garcinia are densly spinulate. In
most taxa, the nexine is < 1 mm thick, but in many Symphonieae it
is > 1.5 mm thick (Stevens, 2007, and literature herein).
Garcinia volkensii Engl (Plate 14, 21e23). (Clusiaceae-
Guttiferae).
Shape: triangular ~29 mm, suboblate.
Apertures: 3-colporate, syncolpate, colpi recessed, pori incon-
spicuously elliptic tall.
Exine: tectate, strongly verrucate, verrucae of variable size and
height.
Habitus: much branched evergreen tree or shrub (2-) 4e20 m
tall.
Habitat: evergreen forest; (30-) 960e2400 m. At Kilimanjaro
mainly in lower to middle montane Ocotea forests; 1870e2540 m.
Distribution: K 3, 4, 6, 7; T 2, 3, 5e7; Zaire, Rwanda, Burundi,
Mozambique, Malawi and Zambia. At Kilimanjaro widespread.
Pollination: most Garcinia species are pollinated by insects, such
as social bees (Pangsuban et al., 2007).
Hypericum revolutum Vahl ssp. keniense (Schweinf.) N. Robson
(Plate 14, 24e26).
Shape: subprolate, trilobate to circular, ~18  23 mm.
Apertures: 3-colporate, colpi (4/5) open with thin margo and
acute end, pori elliptic broad.
Exine: tectate, psilate to finely foveolate.
Habitus: much-branched evergreen shrub or small tree, (0.3-)
1e12 m high.
Habitat: widespread, at forest margins, in fringing and second-
ary forest, scrub, savanna, grassland, on dry or marshy ground;
forest ravines; 900e4000 m. At Kilimanjaro mainly in upper
montane Podocarpus, Juniperus, Hagenia and Erica excelsa forest,
1350e3500 m.
Distribution: T2; in south-western Arabia and the mountains of
East Africa from Eritrea and Ethiopia to Cape Prov. Cameroons,
Fernando Po, Madagascar, Comoro Is., Re union. At Kilimanjaro
widespread.
Pollination: visited by less-specialized insects, of which the
Syrphidae (Diptera) are the most common to Hypericum (Robson,
1981, p. 191).
5.2.39. Icacinaceae
Trees, shrubs or climbers. About 57 genera and 400 species with
a subtropical to tropical and pantropical distribution including
South Africa and Eastern Australia (Peng and Howards, 2008).
5.2.39.1. Pollination. Insects (Willemstein, 1987).
5.2.39.2. Pollen morphology. An eurypalynous family with colpate,
colporidate, colporate, porate, porate, forate, or nonaperturate
grains, ranging in shape from oblate to subprolate, and in size from
 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
15 to slightly >60 mm (Erdtman, 1952).
Apodytes dimidiata E. Mey. ex Arn (Plate 14, 27e28).
Shape: triangular ~24  28 mm, oblate.
Apertures: 3-colporate, short (1/5) and straight colpi with
margo, circular pori with vestibulum.
Exine: reticulate, small meshed reticulum.
Habitus: trees or much branched shrubs up to 25 m high.
Habitat: widespread in lowland and upland rain-forest and
forest patches generally; 1000e2500 m. At Kilimanjaro in sub-
montane Croton-Calodendrum forests, lower and middle montane
Cassipourea-Ocotea forests; 1430e2550 m.
Distribution: U 1, 4; K 3e5; T 1, 2, 4e8; DR Congo Republic and
southwards into Angola and South Africa. At Kilimanjaro scattered.
Pollination: insects, no further details known.
5.2.40. Iridaceae
Perennial evergreen or deciduous herbs, occasionally shrubs
with anomalous secondary growth (Klattia, Nivenia, Witsenia),
rarely annuals (Sisyrinchium spp.), or an achlorophyllous sapro-
phyte (Geosiris). A family comprising ca. 70 genera and 1750 ssp.,
cosmopolitan in distribution, but most abundant and diversified in
southern Africa (Goldblatt et al., 1998).
5.2.40.1. Pollination. Pollination ecology of Iridaceae is very diverse
(e.g. Vogel, 1954). Pollination by bees is ancestral. Long-tongued fly
pollination is particularly common in southern Africa, especially in
genera such as Babiana, Lapeirousia and Gladiolus, but it also occurs
in species of Geissorhiza, Hesperantha, Romulea, Tritonia and Wat-
sonia (Goldblatt et al., 1995). Pollination by hawk moths (Sphingi-
dae) is recorded for H. flava. Savannosiphon and species of Gladiolus.
Sunbird pollination is best developed in Gladiolus and Watsonia,
where it has evolved independently in several lineages in southern
and, in case of Gladiolus, tropical Africa as well.
5.2.40.2. Pollen morphology. Pollen grains are normally shed singly,
and are ellipsoid to spherical (Goldblatt et al., 1998). Grains usually
monosulcate (less often 2-colpate, spirapertur(oid)ate, or non-
aperturate). Longest diameter about 35e120 mm. Sexine as a rule
thicker (sometimes much thicker) than nexine, reticulate, retipilate
or tectum perforatum (Erdtman, 1952).
Gladiolus watsonioides Baker (Plate 14, 29e30).
Shape: prolate ~53  78 mm, circular.
Apertures: monosulcate, sulcus (3/5) open with granulate and
often fissured margins and obtuse ends.
Exine: tectate, collumelate, perforate, microechinate.
Habitus: plants 0.55e1 m high or more. Corm 1.5e2 cm in
diameter.
Habitat: open mountain slopes in the heath zone above the
forest, sometimes in lava rubble, and in glades in Juniperus forest;
2000e4200 m. At Kilimanjaro mainly in upper montane Podo-
carpus, Hagenia and Erica excelsa forests and subalpine ericaceous
bushland; 2450e3690 m.
Distribution: K 3, 4; T 2; not known elsewhere. At Kilimanjaro
scattered.
Pollination: Diptera (Goldblatt et al., 1997; Goldblatt and
Manning, 2006).
5.2.41. Lamiaceae
Trees, shrubs, subshrubs or perennial or annual herbs, rarely
climbers, aromatic or not. The family contains 236 genera and
about 7173 species, almost cosmopolitan, but absent from the
coldest regions of high latitude or altitude (Harley et al., 2004).
5.2.41.1. Pollination. Cross-pollination in Lamiaceae is effected by
animals, of which by far the most diverse group are the insects
355
(Harley et al., 2004).
5.2.41.2. Pollen morphology. Grains of Lamiaceae are single, iso-
polar and, usually, 3 (4) or 6-colpate, oblate to prolate, and longest
axis from 20 to 125 mm. Sexine as thick as nexine or thicker, usually
reticulate. Colpi are usually long with acute apices and provided
with granulate membranes (Erdtman, 1943, 1952).
Clerodendron johnstonii Oliv (Plate 14, 31e33).
Shape: circular ~57 mm, subprolate to spheroidal ~57  63 mm.
Apertures: 3-colpate, straight colpi (2/3 to ¾) open with irreg-
ular margins and obtuse ends.
Exine: tectate, perforate, (micro) echinate.
Habitus: variable; shrub or small tree up to 3e4.5 m with
spreading branches or scrambling shrub 2.5e10.5 m; on Kili-
manjaro a liana up to 20 m.
Habitat: submontane and montane forest and derived thicket,
scrub, cultivations and wasteland, bamboo forest, grassland mar-
gins; 1200e2550 m. At Kilimanjaro in colline and submontane
riverine forests, lower to middle montane Cassipourea and Ocotea
forests as well as in Chagga homegardens; 990e2600 m.
Distribution: U 2, 3; K 1e5, ?6, 7; T 2e4, 6e8; Zaire, Rwanda,
Burundi, Ethiopia, Malawi and NE. Zambia. At Kilimanjaro
widespread.
Pollination: probably various insects (Sakamoto et al., 2012) or
birds (Prasad and Sunojkumar, 2014a).
Leonotis mollissima Gürke (Plate 14, 34e36) SYN L. ocymifolia var.
raineriana.
Shape: (sub)prolate ~34  47 mm, trilobate-circular.
Apertures: 3-colpate, colpi (4/5) narrow (in equatorial view),
recessed with thin margo and acute ends.
Exine: tectate, collumelate, reticulate, heterobrochate.
Habitus: robust, erect, semi-woody shrub, 0.9e5 m tall. Sparsely
branched.
Habitat: margins of montane forest, associated bushland and
grassland, also in ericaceous and bamboo zones, extending down to
Brachystegia woodland and particularly along streams, usually on
well-drained volcanic soils; (50-)600e3700 m. At Kilimanjaro on
colline to submontane roadsides, fallow arable fields and waste
places as well as ericaceous bushland; 1000e2640 m.
Distribution: U 1, 3, 4; K 2e6; T 1e8; DR Congo-Kinshasa,
Rwanda, Burundi, Sudan, Ethiopia, Eritrea, Angola, Zambia,
Malawi, Mozambique, Zimbabwe, Namibia, Swaziland, Lesotho,
South Africa. At Kilimanjaro scattered.
Pollination: by sunbirds, e.g. Nectarinia reichenowi, and by bees
(Gill and Conway, 1979; Vos et al., 1994).
Leucas volkensii Gürke (Plate 14, 37e40).
Shape: subprolate ~25  31 mm, circular to slightly trilobate.
Apertures: 3-colpate, colpi (4/5) narrow with thin margo and
acute ends, granular colpal margins.
Exine: tectate, collumelate, faintly and finely (micro) reticulate.
Habitus: erect herb or shrub with branching stems, 1e3 m high,
often from a woody rootstock.
Habitat: forest (often Hagenia) understorey; 2450e3350 m. At
Kilimanjaro in upper montane Podocarpus, Hagenia and Erica
excelsa forests; 2580e3230 m.
Distribution: T 2; not known elsewhere. At Kilimanjaro wide-
spread in the upper forest belt.
Pollination: not known for this species. Sawflies, wasps, bees
and ants have been observed to pollinate other Leucas species
(Prasad and Sunojkumar, 2014b, 2014c).
Plectranthus alboviolaceus Gürke (Plate 14, 41e42).
Shape: (sub)prolate ~33  45 mm, circular ~40 mm.
Apertures: 6-colpate, colpi (5/6) open in polar view, narrow in
equatorial view.
Exine: tectate, reticulate, lumina smaller in polar area.
356 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386

Habitus: perennial soft-wooded shrub 0.5e3 (4) m tall.
Habitat: evergreen forest, often by paths or rivers;
1500e2800 m. At Kilimanjaro in lower to middle montane Cassi-
pourea, Ocotea forests and riverine forests and forest clearings;
1300e2670 m.
Distribution: K 4, 7; T 1e7; DR Congo, Ethiopia, Zambia, Malawi
and Mozambique. At Kilimanjaro widespread.
Pollination: by bees (Potgieter et al., 2009).
Plectranthus laxiflorus Benth (Plate 14, 43e44 & XV, 1). SYN
P. albus Gürke.
Shape: subprolate to spheroidal ~34  40 mm, circular ~41 mm.
Apertures: 6 colpate, colpi (5/6) open, narrow in equatorial view,
wide in polar view.
Exine: tectate, reticulate, lumina smaller in polar area, muri with
open ends in the equator area, partly very long lumina, lumina
generally larger than in P. alboviolaceus.
Habitus: perennial herb with stems scandent to 4 m.
Habitat: forest margins or clearings, bamboo zone:
1600e2900 m. At Kilimanjaro in submontane riverine and Croton-
Calodendrum forests, lower and middle montane Cassipourea
forests and forest clearings and upper montane Juniperus forests;
1600e2600 m.
Distribution: U 2; K 3, 4; T 2, 3, 6e8; DR Congo, Rwanda, Zambia,
Malawi, Mozambique, Zimbabwe and South Africa. At Kilimanjaro
scattered.
Pollination: by bees (Potgieter et al., 2009).
Plectranthus sylvestris Gürke (Plate 15, 2e4).
Shape: prolate ~42  52 mm, circular ~42 mm.
Apertures: 6 colpate, colpi (5/6) open, narrow in equatorial view,
wide in polar view.
Exine: tectate, ectexine thicker than endexine, exine distinctly
thicker at poles, reticulate with very regular sized lumina, except
for the polar area: here rather reticupilate.
Habitus: soft-wooded shrub or herb, 0.5e3 m tall. Stems erect or
scandent.
Habitat: montane forest, often in bamboo or Hagenia zones;
1700e3300 m. At Kilimanjaro in middle montane Ocotea forests,
upper montane Podocarpus, Hagenia and Erica excelsa forests and
forest clearings; 1950e3090 m.
Distribution: U 2, 3; K 1e6; T 2e4, 6, 7; Nigeria, Cameroon, DR

Plate 15. LAMIACEAE: 1 Plectranthus laxiflorus, 2e4 Plectranthus sylvestris; LAURACEAE: 5e6 Ocotea usambarensis; Linaceae: 7e8 Linum volkensii, LOBELIACEAE (Campanulaceae):
9e11 Cyphia glandulifera, 12e14 Lobelia deckenii, 15e17 Lobelia holstii, LOGANIACEAE (partly Stilbaceae): 18e21 Nuxia congesta, 22e23 Strychnos scheffleri, LORANTHACEAE: 24e25
Agelanthus elegantulus, 26e27 Englerina woodfordioides, 28e30 Plicosepalus curviflorus, 31e32 Tapinanthus brunneus, MALVACEAE: 33e35 Abutilon longicuspe var. longicuspe.
 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
Congo, Rwanda, Ethiopia, Eritrea, Zambia, Malawi and Madagascar.
At Kilimanjaro widespread.
Pollination: by bees (Potgieter et al., 2009).
5.2.42. Lauraceae
Trees or shrubs, mostly evergreen. A pantropical family with a
few temperate members. About 50 genera, number of species
difficult to estimate, probably between 2500 and 3500 (Rohwer,
1993).
5.2.42.1. Pollination. Probably all Lauraceae are obligate out-
breeders, and are insect pollinated (Willemstein, 1987).
5.2.42.2. Pollen morphology. Pollen grains are nonaperturate and
usually more or less spheroidal, with a diameter of (14-)18e40(-70)
mm (Erdtman, 1952). The exine is extremely thin, often seemingly
discontinuous, entirely ectexinous and microechinate (spinules).
The intine is rather thick, and stratified.
Ocotea usambarensis Engl (Plate 15, 5e6).
Shape: spheroidal and circular but often deformed ~ 32 mm,
NOTE: mostly badly preserved.
Apertures: inaperturate.
Exine: very thin, microechinate.
Habitus: tree 3.5e36 (45) m tall, girth (1.25-) 3.75e9.5 m,
unbranched for about 9e15 m.
Habitat: intermediate to montane evergreen wet forest (some-
times dominant in some associations, e.g. Ocotea-Olea); 900e2400
(3000 on Kilimanjaro fide Greenway) m. At Kilimanjaro only on
the southern and eastern slope, mainly in lower to middle montane
Ocotea forests; 1460e2930 m.
Distribution: U 2; K 3, 4, 7; T 2, 3, 6e8; E Zaire, SW Rwanda, N.
Malawi, N. Zambia. At Kilimanjaro widespread.
Pollination: insects; probably thrips as in other Ocotea species
(Danieli-Silva and Varassin, 2013).
5.2.43. Linaceae
Herbs, shrubs, trees, or lianas, sometimes with climbing hooks.
A family of 13 genera with ca. 225 species, mostly from northern
temperate to tropical regions (Dressler et al., 2014).
5.2.43.1. Pollination. Some Linum species have been reported to be
pollinated by bees or flies; self-pollination also occasionally occurs
(Dressler et al., 2014, and literature herein).
5.2.43.2. Pollen morphology. Heterogenous, ±eurypalynous family.
Pollen grains are 3-colpate, 3-colporate, 7e9 colp(orid)ate, poly-
porate, nonaperturate, etc. suboblate to prolate (Erdtman, 1952).
Linum volkensii (Plate 15, 7e8).
Shape: spheroidal ~62  70 mm, trilobate ~75 mm.
Apertures: 3-colpate, colpi long (4/5) and wide with acute ends.
Exine: tectate, exine thick, clavate with polymorphic truncate
clavae.
Habitus: erect, annual or prennial herb up to 0.9 m high.
Habitat: upland grassland, in marshes and by streams;
1300e2750 m. At Kilimanjaro in dry savanna grasslands;
1015e1710 m.
Distribution: U 1e3; K 1e6; T 2e4, 7, 8; S Ethiopia, E DR Congo,
Malawi, Mozambique. At Kilimanjaro scattered.
Pollination: probably insects.
5.2.44. Lobeliaceae (now part of Campanulaceae)
Annual or perennial herbs, subshrubs, shrubs or trees. About 30
genera and some 1000 species, mainly in the tropics and subtropics,
most numerous in the New World (Lobeliaceae: Knox et al., 2008).
357
5.2.44.1. Pollination. Most species are zoophilous. Those with
actinomorphic flowers may be visited by generalized insects,
including bees, flies, wasps, butterflies and settling moths. Orni-
thophily is well documented among Lobelioideae (Lammers, 2009).
5.2.44.2. Pollen morphology. Grains usually (2-) 3 (4) colporate,
prolate to spheroidal, or variously compressed. Sexine thicker than
nexine, pilate to sympilate (Erdtman, 1952). In some Lobelioideae,
the pori may be lacking and the grains are only 3-colpate. Spinules
are common in other subfamilies but lacking in Lobelioideae
(Lammers, 2009).
Cyphia glandulifera Hochst. ex A. Rich (Plate 15, 9e11). (Cam-
panulaceae Cyphioideae).
Shape: subprolate ~31  37 mm, triangular/trilobate.
Apertures: 3-colporate, recessed colpi (4/5) wide with margo
and obtuse ends, constricted at equator. Pori present but unclear in
shape.
Exine: tectate, psilate.
Habitus: erect or somewhat twining herb, 12e60 cm tall. From a
tuber up to ± 5  3 cm. Stem not or sparsely branched. Sessile
leaves, ± in a rosette at the base.
Habitat: grassland and woodland, often in wet depressions;
600e1920 m. At Kilimanjaro in dry colline savanna grasslands;
920e1040 m.
Distribution: K 1e4, 6, 7; T 1, 2, 5; Ethiopia, Somalia. At Kili-
manjaro scattered.
Pollination: insects; maybe pollinated by bees and/or beetles as
in other Cyphia species (Gibson et al., 2012).
Lobelia deckenii (Asch.) Hemsl (Plate 15, 12e14). (Campanula-
ceae Lobelioideae).
Shape: spheroidal, circular to slightly lobate ~36 mm.
Apertures: 3-colporate, colpi (4/5) wide with thin margo and
acute ends, constricted at equator. Colpal area granular. Pori pre-
sent but unclear in shape.
Exine: tectate, scabrate/micropilate.
Habitus: plant 0.8e5 m high.
Habitat: wet moorland, streamsides; 2440e4300 m. At Kili-
manjaro in upper montane Erica excelsa forests, subalpine Erica
trimera forests and ericaceous bushlands and in alpine Helichrysum
srub; 2670e4200 m.
Distribution: T 2; endemic to Kilimanjaro. At Kilimanjaro
scattered.
Pollination: by two sunbird species, i.e. Nectarinia johnstonii and
Cercomela sordida (Burd, 1995).
Lobelia holstii Engl (Plate 15, 15e17). (Campanulaceae
Lobelioideae).
Shape: subprolate ~31  39 mm, spheroidal to trilobate.
Apertures: 3-colporate, colpi (4/5) broad with thick margo and
obtuse ends, constricted at equator. Colpal area granular. Pori pre-
sent but unclear in shape.
Exine: tectate, micro(sym)pilate.
Habitus: perennial herb, 0.1e0.5 m tall, ± erect from a decum-
bent base.
Habitat: upland grassland and moor, forest edges, rocky or
disturbed places; 900e3000 (3500) m. At Kilimanjaro in upper
montane Erica excelsa and Hagenia forests as well as in open hab-
itats (dry lower montane Bulbostylis grasslands and subalpine Fes-
tuca obturbans tussock grasslands); 1590e3000 m.
Distribution: K 1e7; T 2, 3, 5e7; Ethiopia, E. Zaire, Rwanda,
Burundi. At Kilimanjaro scattered.
Pollination: not known; maybe birds as documented among
other Lobelia species.
5.2.45. Loganiaceae (today partly Stilbaceae)
Loganiaceae: family of woody vines, shrubs, or trees containing
358 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
13 genera with more than 400 species native primarily to tropical
areas of the world (Mabberley, 2008; APG, 2009).
Stilbaceae: trees, small shrubs or shrublets. A family of seven
genera and 28 species, widespread in Africa, Madagascar and
Mascarene Islands (Linder, 2004).
5.2.45.1. Pollination. Mammals (Roubik et al., 2005). In Stilbaceae
large, red and black flowers are bird pollinated; rest of family un-
known but unlikely that small white flowers are bird pollinated
(Linder, 2004).
5.2.45.2. Pollen morphology. A hetergenous, eurypalynous family.
Pollen grains are colpate, colporate or porate (Erdtman, 1952, 1957).
Despite the fact that the family of Loganiaceae as used by Erdtmann
(Erdtman, 1952) and in the FTEA (1955e2012) does not hold true
anymore today, Punt and Leenhouts (1967) describe several pollen
types found within genera previously included in this family.
Nuxia congesta R.Br. ex Fresen (Plate 15, 18e21). (Stilbaceae).
Shape: trilobate, spheroidal ~12 mm.
Apertures: 3-colporate, long and wide colpi (5/6), constricted at
equator, pori indistinct.
Exine: collumellate, perforate, tectate, psilate, finely scabrate.
Habitus: tree to 25 m tall.
Habitat: upland rain-forest; 1800e2700 m. At Kilimanjaro in
lower to middle montane Cassipourea and Ocotea forests, however
mainly in upper montane Juniperus forests; 1870e2660 m.
Distribution: U 1e3; K 1, 3, 4, 6; T 2e4, 6, 7; DR Congo, Sudan,
Ethiopia, Malawi, Southern Zimbabwe, and probably farther to the
west and south. At Kilimanjaro widespread in the upper forest belt.
Pollination: self-pollinated as well as by bees in search of nectar
and pollen (Von Breitenbach, 1965). Numerous other insects also
effect pollination and their presence in turn attracts many insec-
tivorous birds.
Strychnos scheffleri Gilg & Busse (Plate 15, 22e23).
Shape: suboblate to oblate, triangular ~42 mm.
Apertures: 3-colporate, colpi (4/5) open with acute ends; pori
inconspicuously lolongate.
Exine: tectate, (micro) reticulate.
Habitus: woody climber up to 25 m high.
Habitat: lowland rain-forest and its secondary growth;
300e1000 m. At Kilimanjaro only in the lower forest types, in
colline savanna and riverine forests and in submontane riverine
and Croton-Calodendrum forests; 960e1310 m.
Distribution: K 7; T 2, 3, 4, 8; possibly Z; not known elsewhere.
At Kilimanjaro scattered.
Pollination: not know for this species. Strychnos species are
probably pollinated by butterflies and moths (Croat, 1978).
5.2.46. Loranthaceae
Evergreen, perennial plants, parasitic, mostly on the branches
(rarely roots) of woody dicotyledons, infrequently on Gymno-
sperms, shrub-like. About 950 species in 77 genera. Mostly tropical
and subtropical, rarely reaching the temperate zones, with several
centers of biodiversity: tropical America, Africa, tropical Asia, and
New Zealand and Australia (Kuijt et al., 2015).
5.2.46.1. Pollen morphology. (2-) 3 (4) colpate, less frequently
colpor(id)ate, peroblate to subprolate. Exine stratification often
obscure. Grains in some species provided with blunt echinae. Amb
usually triangular, in some species distinctly three-armed. Aper-
tures tenuimarginate. Grains sometimes syncolpate (Erdtman,
1952).
Agelanthus elegantulus (Engl.) Polhill & Wiens (Plate 15, 24e25).
Shape: suboblate to oblate, distinctly three-armed Ø ~38 mm,
arm length ~12 mm.
Apertures: 3-colpate, syncolpate, narrow and short colpi.
Exine: tectate, endexine reduced, microechinate.
Habitus: parasitic epiphyte, small shrub, spreading and rather
compact, to 1 m tall.
Habitat: montane forest and upland dry evergreen forest, on a
wide variety of hosts; 1500e2400 m. At Kilimanjaro in lower and
middle montane Ocotea forests and upper montane Juniperus for-
ests; 1640e2650 m.
Distribution: K 4; T 2, 3, 6; not known elsewhere. At Kilimanjaro
scattered.
Pollination: bird-pollinated (Kuijt et al., 2015).
Englerina woodfordioides Balle (Plate 15, 26e27).
Shape: suboblate to oblate, distinctly three-armed Ø ~ 43 mm,
arm length ~20 mm.
Apertures: 3-colpate, syncolpate, narrow colpi which are
slightly recessed.
Exine: tectate, endexine reduced, scabrate.
Habitus: parasitic epiphyte, shrub with spreading to drooping
stems to 1 m or so.
Habitat: montane or riverine forest and associated bushland or
wooded grassland nearby, on many hosts; 1350e3000 m. At Kili-
manjaro in submontane Croton-Calodendrum forests and lower
montane Cassipourea and Ocotea forests; 1460e1680 m.
Distribution: U 1e3; K 1e6; T 2, 3; E. Zaire, Rwanda, Burundi,
Ethiopia. At Kilimanjaro scattered.
Pollination: bird-pollinated (Kuijt et al., 2015).
Plicosepalus curviflorus (Benth. ex Oliv.) Tiegh (Plate 15, 28e30).
Shape: triangular concave, longest axis ~33 mm, oblate.
Apertures: 3-colporate, syncolpate, narrow colpi with distinct
margo, circular pori, pore area thickened.
Exine: tectate, reduced endexine, microechinulate.
Habitus: woody parasitic epiphyte.
Habitat: deciduous bushland and Acacia woodland, almost al-
ways on Acacia, rarely on Albizia; 50e2300 m. At Kilimanjaro in dry
colline savanna forests and woodlands, as well as in Chagga
homegardens; 1040e1600 m.
Distribution: U 1, 3; K 1e7; T 1e3, 7; E Zaire, Somalia, Ethiopia,
along the Red Sea coast to southernmost Egypt, Middle East,
Arabian Peninsula. At Kilimanjaro scattered.
Pollination: bird-pollinated; probably sunbirds like in other
Plicosepalus sp. (Feehan, 1985).
Tapinanthus brunneus (Engl.) Danser (Plate 15, 31e32) SYN
Agelanthes brunneus (FTEA).
Shape: triangular concave ~37 mm, arms ‘rectangular’ in shape,
oblate.
Apertures: 3-colpate, syncolpate, narrow colpi without margo.
Exine: tectate, reduced endexine, scabrate.
Habitus: stems to 1 m or so.
Habitat: evergreen forest, swamp and riverine forest;
1000e1800 m. At Kilimanjaro in lower montane Ocotea forests,
1700e1800 m.
Distribution: U 1, 2, 4; K 5; T 1, 2; westwards to Senegal and
Angola. At Kilimanjaro scattered.
Pollination: not known, maybe sunbird like other Tapinanthus
species (Holm-Nielsen et al., 1989).
5.2.47. Malvaceae
Trees, shrubs or herbs, exceptionally climbers. A cosmopolitan
family of 243 genera and probably more than 4300 species (Bayer
and Kubitzki, 2003). Latest taxonomic results reveal 9 subfamilies
within this family of which only species of 8 subfamilies occur in
tropical East Africa (Stevens, 2015): Grewioideae (see also Tilia-
ceae) include 25 genera and about 770 species and have a
pantropical (warm temperate) distribution. The subfamily of
Byttnerioideae has 26 genera and about 650 species. The
 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
distribution is pantropical with a focus on South America, and also
Australia. The Sterculiodeae (see also Sterculiaceae) are pantrop-
ical and consist of 12 genera and 430 species. The subfamily of
Dombeyoideae has 21 genera and 381 species wich are centred in
the Old World tropics, Australia, St Helena, esp. Madagascar and
Mascarenes. Brownlowioideae include 8 genera and 68 species
with a tropical, esp. Old World distribution. The Malvoideae is the
largest subfamily with 78 genera and 1670 species; its distribution
ranges from temperate to tropical areas. The Bombacoideae (see
also Bombacaceae) include 16 genera and 120 species with a
tropical distribution especially centred in the Neo- and Paleo-
tropics. Helicterioideae comprise 8e10 genera and about 95 species
with a tropical distribution, esp. SE Asia and W Malasia.
5.2.47.1. Pollination. Floral structures and modes of pollination are
very divers within the Malvaceae (Bayer and Kubitzki, 2003 and
literature herein). In Grewioideae the relatively small-flowered
genera point to bee pollination. Larger flowers within this sub-
family are pollinated by vertebrates which include bats and moths.
Most Bombacoideae have large massive sometimes gigantic flowers
which are pollinated by vertebrates, yet occasionally also by bees. In
Byttnerioideae pollinators are bees, moths and also various flies. In
Tiliodeae pollination is primarily by insects but also by wind. Birds,
bats and various insects pollinate species of Helicterioideae. The
flowers of Brownlowioideae all seem to be melittophilous, but no
reports of pollination have been recorded. Within Sterculiodeae,
flowers are rather specialized and fly-mediated sapromyphily is
frequent. Some Dombeyoideae exhibit secondary pollen presenta-
tion, however pollinators have never been reported (except for
Dombeya which seems to be melittophilous). In Malvoideae, bees
and birds are the main pollinators.
5.2.47.2. Pollen morphology. Roughly, the pollen types of the Mal-
vaceae can divided into a few main types (Bayer and Kubitzki,
2003): the Grewia-type, the Tilia-type, the Helicteres-type, the
Bombax-type and the Malva-type. The Malva-type is globose or
rarely (typical of Malvoideae and Dombeyoideae, and some Bom-
bacoideae, Radyera) somewhat flattened and covered
with ± prominent spines, these may all be of the same or of
different length. Apertures vary from zono-3-colporate to zono-
triporate to zono-spiraporate to pantoporate with up to 300 pori.
(The Bombax-type can be found in Bombacaceae, the Grewia-
type in the section onTiliaceae in which the corresponding species
are also described).
Abutilon longicuspe Hochst. ex A. Rich. var longicuspe (Plate 15,
33e35) (Malvoidea).
Malva-type.
Shape: circular, spheroidal ~75 mm.
Apertures: 3 zonoporate, pori lalongate with thin annulus.
Exine: tectum micro-verrucate to micro-granulate, echinate,
columellae distinct, rod-like, elongated near and beneath the bases
of spines, spines short, pointed with a small basal cushion.
Habitus: shrubby herb to softly woody shrub (even described as
small tree) 1e4.5 (5.5) m tall.
Habitat: open Podocarpus, Juniperus, Ekebergia etc. forest, forest
edges, riverine forest and woodland, hillside thicket;
1200e2400 m. At Kilimanjaro in submontane to lower montane
forest clearings and secondary bushlands; 1800e2200 m.
Distribution: U 1e3, K 1e7; T 1e5, 7; Sudan Eritrea, Ethiopia,
Zambia, Malawi, Mozambique, Zimbabwe, Yemen. At Kilimanjaro
scattered.
Pollination: not known for this species. Probably, insects, birds
or bats like in other Abutilon species (e.g. Buzato et al., 1994; Abid
et al., 2010).
Hibiscus meyeri Harv (Plate 16, 1e2). (Malvoidea).
359
Malva-type.
Shape: circular, spheroidal ~90 mm.
Apertures: polyporate with many small circular pori, ± annulate.
Exine: tectum micro-granular, echinate, columellae rod-like,
elongated near or beneath the bases of spines, spines long
~10 mm, straight, slender, with blunt apices and broader base,
sparsely distributed.
Habitus: erect herb 0.5e2 m tall.
Habitat: grassland and bushland; 550e2000 m. At Kilimanjaro
dry savanna bush- and grasslands; 1000 m.
Distribution: U 1e2; K 1e7; T 1e7; DR Congo, Burundi, Ethiopia,
Somalia, Mozambique, South Africa. At Kilimanjaro scattered.
Pollination: not known for this species. Hibiscus in mostly
pollinated by various insects (Faegri and van der Pijl, 1979).
Hibiscus adoensis Hochst. ex A. Rich. (Plate 16, 3e4) Kosteletzkya
adoensis (Hochst. ex A. Rich.) Mast. (Malvoidea).
Malva-type.
Shape: circular, spheroidal ~100 mm.
Apertures: polyporate with many small circular pori.
Exine: thick, tectum micro verrucate/granulate, echinate, colu-
mellae rod-like, slightly elongated beneath the bases of spines,
spines clubbed ~6e10 mm with ± acute apices.
Habitus: perennial herb or subshrub, climbing, scrambling or
procumbent, 0.5e3 m.
Habitat: clearings, secondary growth and forest edges in
montane gallery forest, grassland thicket and scrub, often near
streams, marshes and lake edges, and adventive into pastures,
gardens and abandoned cultivated ground, in partial shade or full
sun, 1000e2700 m. At Kilimanjaro at lower montane thicked edges,
secondary bushlands and heavily disturbed forests; 1600e1800 m.
Distribution: U 2, 3; K 2e7; T 1e4, 6e8; West Africa from Sierra
Leone to Ethiopia and south to Zimbabwe; Madagascar. At Kili-
manjaro widespread.
Pollination: not known for this species. Probably, insects, birds
or bats like in other Abutilon species (e.g. Buzato et al., 1994; Abid
et al., 2010).
Pavonia elegans Garcke (Plate 16, 5e6) (Malvoidea).
Malva-type.
Shape: circular, spheroidal ~110 mm.
Apertures: polyporate; well defined, evenly placed, circular pori
Ø 4e5 mm.
Exine: tectate, collumelate, echinate, monomorphic spines
~15e20 mm long with subacute tips, columellae rod-like, slightly
elongated beneath the bases of spines, sparsely distributed.
Habitus: shrubby herb or shrub 0.6e1.2 m tall.
Habitat: mixed dry evergreen bushland, Acacia-Terminalia and
Adansonia woodland, Acacia-Commiphora bushland, grassland and
cultivations often on black cotton soil; 13e1550 m. At Kilimanjaro
in dry savanna grasslands; 940e1890 m.
Distribution: K 1, 3, 4, 6, 7; T 2, 3; Ethiopia, Somalia. At Kili-
manjaro scattered.
Pollination: generally Pavonia is pollinated by birds and insects
(Gottsberger, 1986).
Sida ovata Forssk (Plate 16, 7e9). (Malvoidea).
Malva-type.
Shape: circular, spheroidal ~104 mm.
Apertures: 5-porate, pori elongate, often obscure.
Exine: columellae thick and densely crowded, distinctly elon-
gated near and at the bases of spines. tectum rugulose to micro-
verrucate, spines short, pointed almost drop-shaped with basal
cushions, densely distributed.
Habitat: grassland (often overgrazed), grassland in scattered
Acacia, Olea etc.; Acacia-Commiphora-Tarchonanthus-Salvadora
bushland and thicket on gravelly, sandy alluvial, hard pan and black
cotton soil; often in rocky places, wadi edges, also a weed in pasture
360 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
Plate 16. MALVACEAE: 1e2 Hibiscus meyeri, 3e4 Hibiscus adoensis, 5e6 Pavonia elegans, 7e9 Sida ovata; MELIACEAE: 10e12 Lepidotrichilia volkensii, 13e15 Trichilia emetica.
and old cultivations; 5e2350 m. At Kilimanjaro in dry savanna
bush- and grasslands, colline to submontane ruderal vegetation on
fallow arable land, roadsides and Chagga homegardens;
800e1900 m.
Distribution: U 1e3; K 1e4, 6, 7; T 1e7; widespread in drier
areas of tropical Africa and South Africa extending to Egypt, Arabia
and India, Iran and Pakistan. At Kilimanjaro very widespread.
Pollination: various insects (Abid et al., 1994).
5.2.48. Meliaceae
Trees, or shrubs (or suckering shrublets), or herbs (rarely, e.g.
Naregamia). Species 575. About 50 genera. Tropical (mostly), or
sub-tropical (few). Pantropical to subtropical and warm
(Mabberley, 2011).
5.2.48.1. Pollination. Most species appear to be insect pollinated,
the agents possibly being bees, sting-less sweatbees or syrphids
while some species are strongly scented particularly in the evening,
which, with their white flowers, suggests moth pollination
(Mabberley, 2011).
5.2.48.2. Pollen morphology. Pollen grains are nearly always radi-
ally symmetrical, suboblate to subprolate monads (Mabberley et al.,
1995). Rarely are they oblate or prolate and only in some Dysoxylum
are they shed in rhomboidal tetrads. They are 3, 4 or 5-colporate,
the second being the most frequent condition. Sexine as a rule
thinner than nexine; the latter usually considerably thickened at
apertures. Pori mostly distinct, circular e lalongate (Erdtman,
1952).
Lepidotrichilia volkensii (Gürke) J.-F. Leroy ex Styles & F. White
(Plate 16, 10e12).
Shape: circular to quadrangular ~21 mm, suboblate to spheroidal.
Apertures: 4e5-colporate, short (1/2) narrow colpi, elliptic
broad/lalongate pori with annulus.
Exine: tectate, exine relatively thick, scabrate.
Habitus: small or medium-sized evergreen tree 5e20 m tall.
Habitat: in montane and mid-altitude forest; 1550e2600 m. At
Kilimanjaro mainly in submontane riverine and Croton-Caloden-
drum forests, lower montane Cassipourea and Ocotea forests;
1280e2560 m.
Distribution: U 1e4; K 1, 3e7; T 2e4, 6e8; Ethiopia, Sudan, E
Zaire, Rwanda, Burundi, Malawi. At Kilimanjaro widespread.
Pollination: insects.
Trichilia emetica Vahl (Plate 16, 13e15).
Shape: subprolate, slightly trilobate/triangular ~ 27 mm.
Apertures: 3-colporate, short slightly opened colpi, pori
 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386 361

lalongate without distinct annulus.
Exine: tectate, psilate.
Habitus: evergreen or semi-evergreen tree, usually 8e20 (25)
m tall.
Habitat: in coastal forest, drier types of riparian forest and ri-
parian woodland, more rarely in rocky outcrops or in wooded
grassland; 10e1300 m. At Kilimanjaro in dry colline savanna and
riverine forests, submontane riverine and Croton-Calodendrum
forests and lower montane gorge forests as well as in Chagga
homegardens; 780e1490 m.
Distribution: U 1e3; K 1, 3e7; T 1e8; Z; widespread in Africa but
absent from the Guineo-DR Congolian region, from Senegal to the
Red Sea and southwards through East and Central Africa to the
Caprivi Strip, Botswana and South Africa, also in the Arabian
Peninsula. At Kilimanjaro widespread in the lower slopes.
Pollination: probably Syrphidae like in other Trichilia species
(Morellato, 2004).
Turraea holstii Gürke (Plate 17, 1e2).
Shape: suboblate, triangular ~53 mm.
Aperture: 3-colporate, very short and narrow colpi, colpal area
granular, pori inconspicuous.
Exine: tectate, collumelate, scabrate, granulate.
Habitus: small tree up to 18 m tall, more rarely a shrub, some-
times scrambling.
Habitat: montane and mid-altitude forest, both in understorey
and at edges, sometimes on stream banks or in abandoned culti-
vation; (700-) 950e2500 m. At Kilimanjaro in colline riverine for-
ests, submontane riverine and Croton-Calodendrum forests, lower
montane riverine and Cassipourea forests; 780e2330 m.
Distribution: U 1; K 2e7; T 2, 3, 5e8; Zaire, Sudan, Ethiopia,
Somalia, Malawi, Arabian Peninsula. At Kilimanjaro widespread,
mainly on the northern and western slope.
Pollination: not known for this species; probably insects like
other Turraea species.

Plate 17. MELIACEAE: 1e2 Turraea holstii, Menispermaceae: 3e5 Stephania abyssinica, MIMOSACEAE: 6e7 Acacia polycantha var. camplyacantha, 8e9 Albizia schimperiana var.
amaniensis, 10e12 Mimosa invisa, MONIMIACEAE: 13e14 Xymalos monospora, MYRSINACEAE (PRIMULACEAE): 15e17 Rapanea melanophloeos, MYRTACEAE: 18e19 Syzygium gui-
neense ssp. afromontanum, OLEACEAE: 20e22 Jasminum abyssinicum, 23e24 Olea capensis ssp. hochstetteri, OXALIDACEAE: 25e27 Oxalis latifolia, PIPERACEAE: 28e29 Peperomia
blanda, 30e31 Piper capense var. capense, PLANTAGINACEAE: 32e34 Plantago palmata, PLUMBAGINACEAE: 35e37 Plumbago dawei, POACEAE: 38e39 Cynodon dactylon.
362 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
5.2.49. Menispermaceae
Twining or rarely erect shrubs or small trees. Pantropical, about
71 genera with 450 species, usually confined to the tropical low-
lands (Kessler, 1993).
5.2.49.1. Pollination. Almost nothing is known about the pollina-
tion of Menispermaceae. Direct observations have never been made
to date and most information from the literature is based on sup-
position (Kessler, 1993). Endress (1996) suggests unspecialized bi-
otic pollination.
5.2.49.2. Pollen morphology. Pollen grains usually 3-colporate (oc-
casionally (2-) 3-colpate (-colporidate), 4e5 rug(or)ate, or non-
aperturate) without operculum and costae, suboblate e prolate.
Sexine as thick as nexine or more frequently thicker, usually per-
reticulate with simplibaculate muri. Pori sometimes operculate
(Erdtman, 1952; Harley and Ferguson, 1982).
Stephania abyssinica (Quart. -Dill. & A. Rich.) Walp (Plate 17,
3e5).
Shape: circular, spheroidal ~12 mm.
Apertures: 3-porate, pori ± circular with small annulus.
Exine: tectate, sexine much thicker than nexine, coarsley per-
reticulate with sparsely distributed granules in the lumina.
Habitus: twining liana, woody at base.
Habitat: varied ecological requirements, not however pene-
trating into the rain-forest; in grassland or wooded grassland up to
3500 m, preferably in moist shady places, especially on the edges of
rivers and swamps. At Kilimanjaro mainly in lower and middle
montane Cassipourea, Ocotea and riverine forests, in upper
montane Podocarpus, Juniperus and Hagenia forests, also in Chagga
homegardens and in open habitats (lower montane grasslands),
1140e2920 m.
Distribution: U 1e4; K 3e5; T 2, 6, 7; widely spread from French
Guinea east to Ethiopia and south through DR Congo to Angola,
Lesotho, South Africa (Natal). At Kilimanjaro widespread.
Pollination: probably unspecialized biotic pollination (Endress,
1996).
5.2.50. Mimosaceae
Trees, shrubs, lianes or rarely herbs, often prickly or spiny. About
60 genera and 3000 species worldwide. Widespread in tropical and
subtropical regions, especially in arid or semi-arid regions (Nielsen,
1992).
5.2.50.1. Pollination. Anemophilous, hydrophilous, entomophilous,
ornithophilous, or cheiropterophilous (Croat, 1978; Endress, 1993).
5.2.50.2. Pollen morphology. Grains are either shed as monads or
united in larger aggregates ranging from tetrads to polyads made
up of up to 32 grains. Sorsa (1969) subdivided pollen grains into five
groups: I) 3-colporate monads, II) large, bisymmetric octads with
pointed peripheral grains, III) radially symmetric, flattened, 12e16
grained polyads, occasionally in tetrads, octads or monads; paraiso-
or heteropolar, pyramidal grains with flattened to convex distal
face; grains generally poroid or rarely 3-colporate, exine verrucose
or gemmate, or sometimes smooth to perforate, IV) grains in tet-
rads, octads or polyads, grains are pyramidal or ± spherical and
small (10e15 mm), exine smooth or granulate to verrucose, V)
grains are radially symmetric, flattened and in 16 (12, 24, or 32)-
celled polyads, grains are (4) 6 (or 8) porate, pyramidal with flat-
tened or slightly convex distal face; exine may be smooth or
foveolate to fossulate.
Acacia polyacantha subsp. camplyacantha (Hochst. ex A. Rich.)
Brenan (Plate 17, 6e7).
Shape: polyads (Ø ~38 mm) of (8 or) 16 grains, single grains
(~11 mm) pyramidal with pointed proximal face.
Apertures: porate?, not discernible in this species.
Exine: pertectate, psilate.
Habit: tree up to 21 m high.
Habitat: wooded grassland, deciduous woodland and bushland,
riverine and ground-water forest; locally common to dominant;
usually gregarious along rivers and in rich, alluvial valleys with
Acacia albida, Kigelia and Ficus sycomorus; 0e1830 m. At Kili-
manjaro in Acacia xanthophloea savanna ground-water forests and
secondary regrowth, 800e1000 m.
Distribution: U 1e4; K 2, 4e7; T 1e8; widespread in tropical
Africa from Gambia to Eritrea in the north and to the Transvaal in
the south. At Kilimanjaro widespread at the foothills.
Pollination: not known for this species, Acacia is generally
pollinated by pollen-foraging insects, bats and birds (Tybirk, 1993;
Stone et al., 2003).
Albizia schimperiana Oliv. var. amaniensis (Baker f.) Brenan (Plate
17, 8e9).
Shape: polyads (Ø ~82 mm) with 16 grains, single grains rect-
angular ~21  26 mm.
Apertures: 6e8 porous areas or faint colpi in the connecting
walls; not clearly discernible in this species.
Exine: tectate, thin exine, finely scabrate.
Habitus: tree 5e23 (30) m high.
Habitat: upland dry evergreen forest, upland rain-forest and
evergreen bushland, riverine forest; 1130e2130 m. At Kilimanjaro
mainly in submontane riverine and Croton-Calodendrum forests, in
lower montane Cassipourea and gorge forests and widely planted
and as a remnant of the former forest cover in Chagga home-
gardens; 930e1770 m.
Distribution: U 1; K 4, 6; T 2, 3, 5, 7; Sudan, Ethiopia, Somalia,
southwards to Portuguese East Africa and Southern Zimbabwe. At
Kilimanjaro widespread.
Pollination: not known.
Mimosa invisa Mart. ex Colla (Plate 17, 10e12).
Shape: tetrade Ø ~25 mm, single grains elliptical ~12 mm.
Apertures: 4-porate, pori with thin annulus.
Exine: tectate, thin exine (<1 mm), finely scabrate.
Habit: shrub up to about 1 m high, often scandent or prostrate,
with long whip-like stems.
Habitat: unknown, an introduced weed; 1310 m. At Kilimanjaro
in colline riverine forests, mainly in Chagga homegardens and
roadsides; 820e1610 m.
Distribution: T 2; native to tropical America, introduced here
and there in the Old World. At Kilimanjaro scattered.
Pollination: Trigona, small bee (Kato et al., 2008). Possibly other
agents as well.
5.2.51. Monimiaceae
Evergreen shrubs or trees, rarely lianas. 34 genera and ca 440
species mainly in the warmer parts of the southern hemisphere
(Philipson, 1993).
5.2.51.1. Pollination. Insect and wind pollinated (Philipson, 1993).
5.2.51.2. Pollen morphology. Grains usually nonaperturate, 2 (3)
sulcate or oligocolpate, tenui-exinous (Erdtman, 1952). The wall is
intectate or tectate and maybe imperforate, perforate or collume-
late; endexine, ectexine and foot-layer may be present or absend.
The surface sculpturing is granular, wrinkled or echinate to varying
degrees (Foreman, 1983).
Xymalos monospora (Harv.) Baill. ex Warb (Plate 17, 13e14).
Shape: (sub)prolate ~27  38 mm, circular to elliptic.
Apertures: inaperturate.
Exine: extremely thin exine, sexine thicker than nexine, clavate
 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
with clavae of different sizes.
Habitus: evergreen shrub or small to medium-sized tree, 6e20
(27) m tall.
Habitat: lowland and upland rain-forest, often a co-dominant in
forests on isolated mountain-tops in dry country; 900e2700 m. At
Kilimanjaro in lower and middle montane Cassipourea, Ocotea and
riverine forests, and in upper montane Podocarpus, Juniperus and
Erica excelsa forests; 1580e2870 m.
Distribution: U 1e4; K 1, ?2, 3e7; T 1e4, 6, 7; eastern Africa from
the Sudan Republic and eastern DR Congo to South Africa, also
Cameroon Highlands and Fernando Po. At Kilimanjaro widespread
in the whole forest belt.
Pollination: wind- or insect-pollinated, but needs further
investigation (Philipson, 1993; Jordaan and Loetter, 2012).
5.2.52. Myrsinaceae (today Primulaceae Myrsinoideae)
Perennial or sometimes annual herbs, subshrubs, shrubs, trees
or lianas. An almost cosmopolitan family of 49 genera and about
1500 species (Ståhl and Anderberg, 2004).
5.2.52.1. Pollination. Most Myrsinaceae appear to be pollinated by
insects, notably bees and flies (Ståhl and Anderberg, 2004). How-
ever, wind pollination has been observed in Rapanea/Myrsine
(Otegui and Cocucci, 1999).
5.2.52.2. Pollen morphology. Pollen grains of Myrsinaceae are
oblate-spheroidal to almost spherical, and 10e45 mm in diameter.
Sexine as thick as nexine or slightly thicker (Erdtman, 1952). Most
taxa have 3-colporate pollen but 4-colporate grains have been re-
ported from Hawaiian Lysiamchia (subg. Lysimachiopsis), Myrsine,
and Rapanea (Ståhl and Anderberg, 2004).
Rapanea melanophloeos (L.) Mez (Plate 17, 15e17) SYN Myrsine
melanophloeos.
Shape: subprolate to spheroidal ~20  22 mm, circular, slightly
trilobate.
Apertures: 3-colporate, short (1/3) slightly open colpi with
irregular margins, circular pori indistinct.
Exine: collumellate, tectate, psilate, scabrate.
Habitus: evergreen tree, or occasionally a shrub 1.8e4.5 m tall.
Very small buttresses and a fluted trunk.
Habitat: upland forest, riverine and swamp forest, open wood-
land, thickets, scrub near streams and in upland grassland on white
sandy or peaty or volcanic soil; (5-)900e3750 m. At Kilimanjaro in
lower and middle montane Cassipourea, Ocotea and riverine forests,
and in upper montane Podocarpus, Juniperus, Hagenia and Erica
excelsa forests; 1530e3240 m.
Distribution: U 1e4; K 2e7; T 2e8; Z; tropical Africa from
Nigeria, Fernando Po, and Cameroon to Ethiopia, south to Angola,
and in South Africa. At Kilimanjaro widespread in the whole forest
belt.
Pollination: maybe wind pollinated like other Myrsine sp.
(Otegui and Cocucci, 1999).
5.2.53. Myrtaceae
Trees and shrubs. Halophytic to xerophytic. Predominantly
tropical to southern temperate distribution but with relatively low
representation in the African region. The family includes around
142 genera and 5500 species (Wilson, 2011).
5.2.53.1. Pollination. Outbreeding is probably widespread,
although both self-compatible and self incompatible species exist.
Bee-pollination with pollen as the reward is a common pollination
system. Bird- and mammal pollination is also known to occur in
Syzygium. Much pollination is generalized or unspecialized and
may be effected by a range of animals, like beetles (observed on
363
Angophora), nectar-feeding fruit bats (observed on Syncarpia) and
even lizards (recorded on Metrosideros) (Wilson, 2011).
5.2.53.2. Pollen morphology. Pollen grains in Myrtaceae are rela-
tively uniformly 3-colporate, radially symmetrical, mainly oblate,
and triangular with straight or curved sides (Gadek and Martin,
1982; Patel et al., 1984). Pike (1956) divided the pollen types into
three categories: (1) longicolpate, (2) syn- or parasyncolpate, and
(3) brevi- or brevissimicolpate.
Syzygium guineense subsp. afromontanum F. White (Plate 17,
18e19).
Shape: oblate, (concave) triangular, ~21 mm.
Apertures: 3-colporate, syncolpate, costae, lalongate pori with
vestibulum.
Exine: tectate, psilate, scabrate, triangular island of thick tectum
at the pole of the grain.
Habit: trees, shrubs or pyrophytic subshrubs 0.2e30 m tall.
Habitat: evergreen forest slopes, escarpment crests, streamsides
but not means always riverine, forest edges also in Acacia, Com-
bretum, Brachystegia woodland; (450-) 900e2250 m. At Kili-
manjaro in submontane riverine and Croton-Calodendrum forests,
lower and middle montane Cassipourea, Ocotea and gorge forests;
1170e2420 m.
Distribution: U 1e3; K 1/2, 3e5, 7; T 2e8; from Sudan to Congo
south to Angola, Mozambique and Zimbabwe widespread
throughout tropical Africa from Senegal to Somalia to South Africa,
Sudan to DR Congo-Kinshasa to Angola, Cameroon, Gabon, Central
Africa, Mozambique, Zimbabwe, Zambia, Saudi Arabia, Yemen. At
Kilimanjaro widespread in the lower half of the forest belt.
Pollination: by honeybees, i.e. Apis mellifera adansonii (Wubie
et al., 2014) and other insects.
5.2.54. Oleaceae
Trees, shrubs or woody climbers. A world-wide family of 25
genera and about 600 species (Green, 2004).
5.2.54.1. Pollination. Entomophilous in most Oleaceae. Cross-
pollination is favoured and some genera are noted for being het-
erostylic. The hypocrateriform corollas of several genera, e.g. Sy-
ringa and Jasminum, provide a flat platform on which a long-
tongued insect can land (Green, 2004).
5.2.54.2. Pollen morphology. Pollen grains are usually 3 (4) col-
pate to colpor(id)ate, oblate to prolate with a reticulate exine.
Sexine mostly thicker than nexine (Erdtman, 1952). While there is
some variation in the sizes of the reticulum and the micromor-
phology of the muri, there is an overall uniformity of structure in
the pollen grains of the whole family.
Jasminum abyssinicum Hochst. ex DC (Plate 17, 20e22).
Shape: spheroidal to subprolate ~42 mm, tetralobate ~38 mm.
Apertures: 4-colpate, colpi (3/5) narrow with acute ends.
Exine: tectate, sexine much thicker than nexine, reticulate,
heterobrochate.
Habitus: generally, a climbing or a scrambling shrub.
Habitat: forest undergrowth and margins, hillsides, bushland,
often near streams; 690e2700 m. At Kilimanjaro in submontane
riverine and Croton-Calodendrum forests and lower montane Cas-
sipourea and gorge forests; 1280e2210 m.
Distribution: U 2e4; K 3e6; T 1, 2, 7; Ethiopia, DR Congo. At
Kilimanjaro scattered in the lower forest belt, mainly on the
northern and western slopes.
Pollination: long-tongued insects (Green, 2004).
Olea capensis subsp. hochstetteri (Baker) Friis & P. S. Green (Plate
17, 23e24).
Shape: trilobate, spheroidal ~23 mm.
364 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
Apertures: 3-colpate, colpi long and very narrow without
distinct margins.
Exine: tectate, thick exine, coarsely reticulate.
Habitus: often a bushy shrub or a small to medium sized tree up
to 10 m in height, but it may be much larger, occasionally reaching
40 m.
Habitat: occurring in bush, littoral scrub and evergreen forest
where temperatures are relatively low and, apart from diurnal
fluctuations, fairly constant temperatures throughout the year. At
Kilimanjaro in submontane riverine and Croton-Calodendrum for-
ests and lower montane Cassipourea and gorge forests, sometimes
also in montane Juniperus forests; 1170e2490 m.
Distribution: occurs almost throughout Africa south of the
Sahara Desert from Sudan and Ethiopia to the southern extremity
of the continent and west to the Islands of the Gulf of Guinea and to
Sierra Leone (Orwa et al., 2009). At Kilimanjaro widespread, but
always in few numbers in the lower forest belt.
Pollination: pollination ecology remains undocumented
although the morphology of the flowers point to moths and small
bees as possible pollinators (Tsingalia, 2010).
5.2.55. Oxalidaceae
Perennial, rarely annual herbs, sometimes succulent, often with
underground storaging bulbs, tubers or rape-like roots, or shrubs,
small trees, or sometimes vines. A family comprising five genera
and about 880 species, widespread in tropical and temperate zones
(Cocucci, 2004).
5.2.55.1. Pollination. Bee pollination prevails but other insects are
possible pollinators and self-pollination exists (Cocucci, 2004).
5.2.55.2. Pollen morphology. Pollen grains are 3-colpate or tricol-
poridate; some Oxalis are 3-colporate and exceptionally pan-
tocolp(or)ate or 4-colp(or)ate, the exine is finely reticulate. The
shape is oblate to spheroidal (Erdtman, 1952).
Oxalis latifolia Kunth (Plate 17, 25e27).
Shape: subprolate/spheroidal ~30  35 mm, circular/trilobate
~33 mm.
Apertures: 3-colpate, colpi long (5/6) and recessed, wider at
equator and with acute ends. Colpi often appear granular inside.
Exine: tectate, reticulate, homobrochate.
Habit: acaulescent herb, with leaves and peduncles arising
directly from the bulb.
Habitat: cultivated ground and waste places, usually in moist
and well-shaded parts; sea-leavel to 2520 m. At Kilimanjaro in
Chagga homegardens; 960e1770 m.
Distribution: U 2e4; K 3e5, 7; T 1e3, 6; DR Congo, Sudan,
Ethiopia, Zambia, Zimbabwe, Mozambique, South Africa; native of
America, introduced to Africa. At Kilimanjaro in the Coffee-banana
belt widespread.
Pollination: insects, no further details available.
5.2.56. Piperaceae
Trees, shrubs, lianas, herbs, sometimes epiphytic, often aro-
matic, with spherical oil cells. A tropical and subtropical family of
five genera and ca. 3000 species (Tebbs, 1993).
5.2.56.1. Pollination. Wind- or insect pollinated (Figueiredo and
Sazima, 2000).
5.2.56.2. Pollen morphology. Pollen grains are globose, tectate,
binucleate, monosulcate or, in Peperomia, inaperturate. The tectum
is often minutely spinulose (Erdtman, 1952).
Peperomia blanda (Jacq.) Kunth (Plate 17, 28e29).
Shape: circular, spheroidal ~11 mm.
Apertures: inaperturate.
Exine: tectate, very thin exine, verrucate or minutely echinate.
Habitus: erect succulent rhizomatous plant, 0.1e0.5 (0.6) m
tall, sometimes epiphytic.
Habitat: bare but often shady rocky places, e.g. granite outcrops,
inselbergs, gullies, evergreen scrub and ‘dry forest’, also margin of
standing water and springs, often with Aloe and Aeollanthus, or in
riverine thicket, rarely an epiphyte, e.g. on Diospyros; 250e1800 m.
At Kilimanjaro in colline e submontane riverine forests;
1080e1220 m.
Distribution: U 2, 3; K 4e7; T 1e3, 6, 7; from Yemen to South
Africa west to Zaire, Madagascar, Mascarene Is., India, Burma, Indo-
China, Taiwan, Malesia, Micronesia, Polynesia, New Caledonia,
Australia, USA (Florida) to S. South America. At Kilimanjaro rare.
Pollination: although the floral morphology of Peperomia spe-
cies suggests wind- and/or insect pollination, most of the species
studied exhibit autogamy and perhaps agamospermy as the main
method of reproduction (Figueiredo and Sazima, 2007).
Piper capense L. f. var. capense (Plate 17, 30e31).
Shape: circular, spheroidal ~11 mm.
Apertures: monosulcate, sulcus mostly wide.
Exine: tectate, very thin, verrucate or minutely echinate.
Habitus: weakly erect shrub or subshrub or ± herbaceous or
sometimes a straggling liana, 1e5 m tall or long.
Habitat: essentially forest undergrowth in wet places, swampy
forest edges, mixed bamboo forest, also upland scrub and thicket
near streams, grassland and tree clumps, etc.; 650e2500 m
(2700 m Mt. Nyiro fide Kerfoot). At Kilimanjaro in submontane
riverine forests, lower e middle montane Cassipourea, Ocotea and
gorge forests and upper montane Juniperus forests; 1170e2900 m.
Distribution: U 2e4; K 1e7; T 1e4, 6, 8; widespread in Africa
from Sierra Leone to Cameroon, Rio Muni, Bioko, Sao Tome  to Zaire,
Rwanda, Burundi, Sudan, Ethiopia, south to Mozambique,
Zimbabwe, Malawi, Swaziland, South Africa, Comoro Is.,
Madagascar. At Kilimanjaro widespread.
Pollination: not known for this species. Figueiredo and Sazima
(2000) identified ambophily (wind and small insects) as the pri-
mary pollination system for southeast Brazilian Piper species. Self-
pollination is not ruled out either.
5.2.57. Plantaginaceae
Herbs or occasionally small shrubs, terrestrial (to aquatic). The
family is cosmopolitan with one genus and about 270 species.
5.2.57.1. Pollination. Most Plantago species are wind pollinated.
However, insect-pollination is found as well. Pollinators are syrphid
and muscid flies, as well as certain bees and beetles (Schwarzbach,
2004, and literature herein).
5.2.57.2. Pollen morphology. Pollen grains are spheroidal and pro-
vided with (3-) 4e14 colpi or pori. Pollen characters do not vary
substantially in the family (Erdtman, 1952; Schwarzbach, 2004).
Plantago palmata Hook. f (Plate 17, 32e34).
Shape: circular, spheroidal ~24 mm.
Apertures: periporate, < 10 roundish pori, sunken with granu-
late margins.
Exine: tectate, thin collumella, verrucate.
Habitus: perennial herb with short, stout rhizome and
numerous roots.
Habitat: openings and disturbed places in montane forest,
particularly bamboo and Juniperus, often by streams or on boggy
ground, less often in thicket or in plantations; 1170e3150 (-? 3300)
m. At Kilimanjaro in lower e upper montane riverine forests,
trampled ground in Chagga homegardens and forest tracks;
1600e3500 m.
 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
Distribution: U 1e4; K 2e4; T 2, 7; Cameroon, Bioko I., E. DR
Congo, Rwanda, Ethiopia, Zimbabwe. At Kilimanjaro widespread.
Pollination: not known for this species. Predominantly
inbreeding Plantago species ensure self-pollination; in outcrossed
species wind and insects are common pollinators (Sharma et al.,
1993).
5.2.58. Plumbaginaceae
Shrubs, lianas, or herbs. The family consists of 27 genera,
embracing about 650 species, distributed throughout the world,
especially in the Mediterranean and Irano-Turanian regions
(Kubitzki, 1993).
5.2.58.1. Pollination. Bees and small beetles have been observed
pollinating Mediterranean leadworts. Pollination by bees, flies and
small beetles has been observed in the Neotropics (Every, 2009).
5.2.58.2. Pollen morphology. Pollen grains are usually 3-colpate,
rarely 4e5 colpate, and, exceptionally, pantocolpate. Two pollen
types can be distinguished (Erdtman, 1952): the Plumbago type,
characteristic of the Plumbaginoideae, and the Armeria type, char-
acterizing the Staticoideae. The Plumbago type has a well-defined
foot-layer, highly irregular columellae, and a continuous tectum.
Upward extending from the latter is another set of columellae,
designated as verrucae. The Armeria type has a foot layer sup-
porting straight, regular columellae, which are distally fused into an
incomplete tectum or reticulate configuration.
Plumbago dawei Rolfe (Plate 17, 35e37).
Shape: (sub)prolate ~50  70 mm, circular.
Apertures: 3-colpate, colpi (5/6) wide, tapering towards poles
with acute ends.
Exine: tectate, columellate, verrucate.
Habitus: creeping or scandent herb or shrub. Laxly branched,
0.9e5.7 m tall.
Habitat: forest margins, clearings and secondary bushland,
riverine forest; 600e2700 m. At Kilimanjaro in colline riverine
forests; 1170 m.
Distribution: U 2e4; K 1, 7; T 2, 5e7; Ethiopia, Madagascar. At
Kilimanjaro rare.
Pollination: not known; probably various insects like in other
Plumbago species (e.g. Ferrero et al., 2009).
5.2.59. Poaceae
Herbs, or shrubs, or ‘arborescent’, or lianas. Annual, or biennial,
or perennial. This family consists of about 12,000 species in ca. 700
genera, is found on all continents and exhibits remarkable
ecological diversity. Soecies occur in tropical forests, in deserts,
from near the poles to the equator and from sea level to high alti-
tudes (Kellogg, 2015).
5.2.59.1. Pollination. Despite the prevalence of wind-pollination in
this family, a few taxa maybe insect pollinated (e.g. Soderstrom and
Calderon, 1971; Sajo et al., 2012).
5.2.59.2. Pollen morphology. Grass pollen is remarkably uniform in
morphology. Pollen grains are more or less spheroidal/circular with
a single annulate pore (except for the bambusoid genus Pariana
where a pore is absent). On the inside of the annulus, the foot layer
is thickened and layered. The exine is columellate and covered with
small granules; these may form islands (clusters) in some species,
whereas the granules are isolated in others (Liu et al., 2004;
Perveen, 2006; Kellogg, 2015, and literature herein). However,
differences between species and genera are usually subtle.
Cynodon dactylon (L.) Pers (Plate 17, 38e39).
Shape: subprolate, circular ~30e33 mm.
365
Apertures: monoporate, annulate (costa and margo), pore
diameter ~2.4e3.3 mm, annulus width ~1.6e1.7 mm, annulus height
~1.5e1.6 mm, operculate, pore in level with grain outline (ectexine).
Exine: tectate, very thin endexine, columellate (not visibly),
scabrate, microechinate, compactly insular exine pattern.
Habitus: stoloniferous sward-forming perennial, culms slender,
0.08e0.4 m high.
Habitat: roadsides, old farmland, and weedy or trodden places
around habitations. Used as a lawn grass throughout East Africa,
0e2000 m. At Kilimanjaro on colline e submontane grasslands as
well as roadsides, fallow arable land, waste places; 890e1890 m.
Distribution: U 1e4; K 1, 3e7; T 1e7; Z; tropical and warm
temperate regions throughout the world. At Kilimanjaro
widespread.
Pollination: wind.
Dactyloctenium aegyptium (L.) Willd (Plate 18, 1e2).
Shape: spheroidal, circular (often deformed) ~31 mm.
Apertures: monoporate, pore Ø 4 mm annulate, annulus width
~2 mm, non-operculate, pore slightly sunken in.
Exine: tectate, very thin endexine, columellate (not visibly),
scabrate, microechinate, sparsely insular.
Habitus: slender to moderately robust spreading annual, culms
up to 0.7 (1) m high.
Habitat: widespread weed of open situations in grassland and
open woodland, common by roadsides and on waste ground, els-
where from 1 to 2100 m. At Kilimanjaro in savanna and saline
grasslands as well as on roadsides, fallow arable land, waste places;
900e1370 m.
Distribution: U 1e4; K 1e7; T 1e8; Z; P; throughout Africa.
Widely distributed in tropical and warm temperate regions of the
Old World, introduced into America. At Kilimanjaro widespread.
Pollination: wind.
Diheteropogon amplectens (Nees) Clayton (Plate 18, 3e4).
Shape: spheroidal, circular ~33 mm.
Apertures: monoporate, pore Ø ~2 mm, annulate, annulus width
~3.5 mm, annulus height ~2.5 mm, operculate, annulus protruding
distinctly.
Exine: tectate, thin exine, psilate to finely scabrate, regular
granule pattern.
Habitus: perennial with short rhizomes, culms 0.2e3 m high.
Habitat: stony slopes and poor sandy soils in deciduous bush-
land, coastal bushland and wooded grassland; 0e2000 m. At Kili-
manjaro in colline and submontane grasslands; 1000e1500 m.
Distribution: K 4e7; T 1e6, 8; Zaire, Mozambique, Zimbabwe,
South Africa. At Kilimanjaro widespread.
Pollination: wind.
Eleusine jaegeri Pilg (Plate 18, 5e6).
Shape: oblate-spheroidal (often deformed) ~25 mm.
Apertures: monoporate, annulate (thin margo, thick costa),
operculate, pore Ø ~2.5 mm in level with grain outline or slightly
sunken, annulus width ~2.5e3 mm.
Exine: tectate, very thin endexine, columellate (not visibly),
scabrate, sprarsely insular.
Habitus: coarse perennial tussock grass from a short ascending
rhizome, culms 0.4e1.3 m high.
Habitat: upland grassland and clearings in forest and bushland,
locally dominant. 1800e3300 m. At Kilimanjaro in montane forest
clearings and submontane grasslands; 2650 m.
Distribution: U 3; K 3e6; T 2; not known elsewhere. At Kili-
manjaro only on the northern slope, rare.
Pollination: wind.
Elionurus muticus (Spreng.) Kuntze (Plate 18, 7e8).
Shape: spheroidal, circular (often deformed) ~40 mm.
Apertures: monoporate, annulate, pore Ø ~3 mm, non-
operculate, annulus height ~3 mm protruding distinctly, annulus
366 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386

Plate 18. POACEAE: 1e2 Dactyloctenium aegyptium, 3e4 Diheteropogon amplectens, 5e6 Eleusine jaegeri, 7e8 Elionurus muticus, 9e10 Harpachne schimperi, 11e13 Hyperthelia
dissoluta, 14e15 Isachne mauritiana, PODOCARPACEAE: 16e18 Podocarpus falcatus, 19e21 Podocarpus latifolius, POLYGONACEAE: 22e23 Polygonum salicifolium, 24e26 Rumex
bequaertii, Proteaceae: 27e28 Grevillea robusta, 29e30 Protea kilimandscharica, RANUNCULACEAE: 31e32 Clematis brachiata.

width ~3.5 mm.
Exine: tectate, very thin endexine, scabrate, granulate.
Habitus: densely tufted perennial, culms 0.15e1 m high.
Habitat: open deciduous bushland on dry stony soils,
1400e2800 m. At Kilimanjaro in colline and submontane grass-
lands; 990e1500 m.
Distribution: U 1e4; K 3e6; T 1, 2, 4, 5, 7, 8; Yemen, tropical and
South Africa, tropical and subtropical America. At Kilimanjaro
widespread.
Pollination: wind.
Harpachne schimperi Hochst. ex A. Rich (Plate 18, 9e10).
Shape: circular, spheroidal (often deformed) ~ 28 mm.
Apertures: monoporate, pore circular Ø ~3.5 mm, annulus width
~3 mm, only slightly protruding, non-operculate.
Exine: tectate, very thin endexine, coarsely scabrate, sprarsely
insular.
Habitus: densely tufted perennial, culms 0.13e0.52 m high.
Habitat: grassland and open bushland, often on dry sandy or
stony soils, or as a weed of roadsides and waste ground,
500e2600 m. At Kilimanjaro in colline e submontane grasslands;
1180e1620 m.
Distribution: U 1e4; K 1e7; T 1e5, 7, 8; Sudan, Ethiopia, N So-
mali Republic southwards to Zambia. At Kilimanjaro scattered.
Pollination: wind.
Hyperthelia dissoluta (Nees ex Steud.) Clayton (Plate 18, 11e13).
Shape: spheroidal, circular (often deformed) ~37 mm.
Apertures: monoporate, annulate, pore Ø ~3 mm circular,
annulus width ~3 mm, annulus height ~2.5 mm, slightly protruding,
sometimes operculate.
Exine: tectate, very thin endexine, coarsely scabrate to granular.
Habitus: tufted perennial, culms 1e3 m high.
Habitat: common species of deciduous bushland and wooded
grassland, particularly by waysides and in disturbed places;
0e2400 m. At Kilimanjaro in colline savanna grasslands;
1020e1390 m.
Distribution: U 1e4; K 2e7; T 1e8; Z; tropical and South Africa,
 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
Madagascar, introduced to tropical America. At Kilimanjaro
widespread.
Pollination: wind.
Isachne mauritiana Baker (Plate 18, 14e15).
Shape: spheroidal, circular (often deformed) ~ 31.
Apertures: monoporate, pore circular Ø ~2 mm with protruding
annulus, annulus width ~1.5 mm and height ~1.5 mm.
Exine: tectate, very thin endexine, scabrate, negative reticulum
with circular to longate areolae, more or less regular granule
pattern.
Habitus: rambling perennial, culms 0.3e0.6 m high.
Habitat: forest shade, 1000e2600 m. At Kilimanjaro mainly in
lower e middle montane Ocotea forests and clearings;
1120e2460 m.
Distribution: U 2, 4; K 3e5, 7; T 1e3, 6, 7; Cameroon and Zaire to
Zimbabwe, Madagascar and Mauritius, a few records from Nigeria
and Ghana. At Kilimanjaro widespread in the forests of the south-
ern slope.
Pollination: wind.
5.2.60. Podocarpaceae
Slightly resinous, evergreen shrubs or trees. At least 17 genera
and 125 or more species, mostly tropical-subtropical montane,
mostly in the southern hemisphere (Page, 1990).
5.2.60.1. Pollination. Pollen transfer by wind (Tomlinson et al.,
1991).
5.2.60.2. Pollen morphology. Grains bilateral-bisaccate or
radiosymmetric-trisaccate (or tetra-saccate, occasionally more or
less synsaccate), analept, without orbicules. Breadth of corpus
35e50 mm, of sacci about 35e65 mm (Erdtman, 1965).
Podocarpus falcatus (Thunb.) R. Br. ex Mirb (Plate 18, 16e18).
Shape: bilateral, heteropolar, vesiculate, bisaccate; corpus
length 27e33 mm, breadth 23e29 mm, depth Ø 22.5 mm; saccus
length Ø 11.6 mm, saccus breadth Ø 26.5 mm, saccus depth Ø 17.4 mm.
Apertures: inaperturate.
Exine: the corpus is psilate to finely scabrate, the sacci are
reticulate, muri high, lumina of variable shape and size.
Habitus: evergreen tree reaching up to 46 m in height with long,
clean cylindrical trunk (Feleke et al., 2012).
Habitat: in coastal swamp forest, transitional rainforest, dry
evergreen forest, undifferential forest, and Afromontane rainforest.
Occasionally in Afromontane rain forest, but particularly charac-
teristic of undifferentiated Afromontane forest, where it is
frequently one of the dominant species or one of the co-dominant
species (e.g. in Juniperus-Podocarpus forest), often persisting in
relict forest patches (gully forests, church forest. Rich, well-drained
soils are needed for P. falcatus; mainly found on humus-rich sandy
soils (Orwa et al., 2009). At Kilimanjaro mainly in Cassipourea and
Juniperus forests; 1280e2230 m.
Distribution: Burundi, DR Congo, Eritrea, Ethiopia, Kenya,
Lesotho, Malawi, Mozambique, Rwanda, South Africa, Sudan,
Swaziland, Tanzania, Uganda, Zimbabwe, India (exotic) (Orwa et al.,
2009). At Kilimanjaro scattered, only on the western and northern
slope.
Pollination: wind (Negash, 2003).
Podocarpus latifolius (Thunb.) R. Br. ex Mirb (Plate 18, 19e21).
Shape: bilateral, heteropolar, vesiculate, bisaccate; corpus about
22  44  32 mm, longest overall diameter about 78 mm, corpus
length 33e43 mm, breadth 29e37 mm, depth Ø 27.4 mm; saccus
length Ø 14.3 mm, saccus breadth Ø 34.3 mm, saccus depth Ø
22.6 mm.
Apertures: inaperturate.
Exine: the corpus is (coarsely) scabrate to granular, the sacci are
367
(incomplete) reticulate, muri often have open ends, lumina large
and of variable shape and size.
Habitus: tree to 35 m high.
Habitat: upland rain-forest; 900e3150 m. At Kilimanjaro mainly
in lower e middle montane Cassipourea forests, middle montane
Ocotea, upper montane Podocarpus, Hagenia and Juniperus forests;
1600e3200 m.
Distribution: U 1, 2, 4; K 1, 3e5; T 2, 3, 5e7; DR Congo, Sudan,
Zimbabwe, Malawi, Angola, Cameroon. At Kilimanjaro widespread.
Pollination: wind (Geldenhuys, C. J., 1993).
5.2.61. Polygonaceae
Annual or perennial herbs, shrubs, trees, or clambering, climb-
ing, or twining lianas. A family of worldwide distribution,
comprising 43 genera and 1100 species, and occupying a wide
range of habitats (Brandbyge, 1993).
5.2.61.1. Pollination. Anemophilous or entomophilous (Brandbyge,
1993).
5.2.61.2. Pollen morphology. The family is palynologically very
divers and the extensive variation has great taxonomic potential at
all levels, especially for the definition of genera (Nowicke and
Skvarla, 1977, 1982). Pollen grains 3e4 (6) colporate, 4 lox-
ocoplorate, polycolpate, polyporate etc., oblate spheroidal to pro-
late (Erdtman, 1952).
Polygonum salicifolium Brouss. ex Willd (Plate 18, 22e23).
Shape: circular, spheroidal ~53 mm.
Apertures: periporate, circular pori.
Exine: tectate, thick, coarsely reticulate, high muri with
bacculae.
Habitus: an erect or basally decumbent, slender annual, up to
1 m tall.
Habitat: damp places, often growing in water; sea level to
2400 m. At Kilimanjaro in swamps and streamsides from colline e
submontane; 790e1680 m.
Distribution: U 2, 4; K ?3, 4, 5; T 1e8; Z; P; throughout tropical
Africa, also in tropical Asia, Australia, and America, naturalized in
Madagascar. At Kilimanjaro scattered.
Pollination: probably various insects like in other Polygonum
species (Hawkes, 1966; Momose and Inoue, 1993).
Rumex bequaertii De Wild (Plate 18, 24e26).
Shape: circular, spheroidal ~27 mm.
Apertures: 3-colporate, colpi (4/5) very narrow, pori elliptic tall,
wider than colpi, faint annulus.
Exine: tectate, coarsely scabrate.
Habitus: an erect, stout, perennial herb, up to 1.8 m tall.
Habitat: upland grassland and bushland, upland rain-forest,
riverside grassland, particularly in damp places; 690e3700 m. At
Kilimanjaro in submontane e upper montane riverine forests and
streamsides, also in Chagga homegardens; 980e2960 m.
Distribution: U 2e4; K 3e5, ?6; T 2, 3, 6e8; widely spread
through eastern Africa reaching Ethiopia and the Transvaal, also in
the Cameroons and Madagascar. At Kilimanjaro widespread.
Pollination: not known for this species. Rumex species are
generally pollinated either by wind or by insects.
5.2.62. Proteaceae
Perennial shrubs or trees, a family comprising 80 genera and
about 1700 species, distributed mainly in the southern hemisphere,
where it is almost completely restricted to Gondwanic continental
blocks and fragments. It is most diverse in Australia, followed by
southern Africa, South America, New Caledonia, Malesia, South and
East Asia, tropical Africa, Central America, Madagascar, New Zea-
land, Fiji, southern India, Sri Lanka, Vanuatu and Micronesia
368 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
(Weston, 2010).
5.2.62.1. Pollination. Known pollinators in Proteaceae include in-
sects such as bees, beetles, flies, butterflies and moths, birds such as
honeyeaters, sunbirds, sugarbirds and hummingbirds, and mam-
mals, including rodents, small marsupials, elephant shrews and
bats, as in Banksia and Protea (Weston, 2010, and literature herein).
5.2.62.2. Pollen morphology. Pollen grains of Proteaceae are prim-
itively triporate, triangular in polar view, with the pori aligned in
the equatorial plane (Johnson and Briggs, 1975). Variation in exine
structure, especially that of the apertures and sculpturing of the
tectum amd supratectal elements, is extensive but is probably
moderately to highly homoplasious (Weston, 2010).
Grevillea robusta A. Cunn. ex R. Br (Plate 18, 27e28).
Shape: triangular ~57 mm, oblate.
Apertures: 3-porate, pori large Ø 11 mm and membranes pro-
truding like a bubble.
Exine: tectate, surface undulating causing a reticuloid pattern.
Habitus: tree up to 30 m high.
Habitat: sometimes more or less established in native vegeta-
tion near habitation or roadsides; recorded 1500e2000 m, but
probably planted more widely. At Kilimanjaro not naturalized,
planted between 820 and 1770 m.
Distribution: Kenya (Naivasha, S. Kavirondo), Tanzania (Arusha,
Buha, Morogoro, Moshi), widely planted as a street tree, as a shade
tree on tea or coffee plantations, or as a garden ornamental, native
of Australia, now widely cultivated in tropical countries. At Kili-
manjaro widespread.
Pollination: by nectarivorous birds, such as honeyeaters, sun-
birds (Nectariniidae) and sugarbirds (Promeropiidae) and the Cape
white-eye Zosterops pallidus, as well as fruit bats, order Mega-
chiroptera (Kalinganire et al., 2001).
Protea kilimandscharica Engl (Plate 18, 29e30). SYN P. caffra
subsp. kilimandscharica.
Shape: (convex) triangular ~24 mm, oblate.
Apertures: 3-porate, pori circular with vestibulum.
Exine: tectate, thin, thinnest towards apertures, scabrate.
Habitus: shrub 1e3.5 m or more, rarely a tree up to 6 m.
Habitat: in ericaceous scrub, heathland, stream banks, rocky
hillsides, montane grassland and forest margins; 2300e3700 m. At
Kilimanjaro in subalpine ericaceous bushland; 2930e3500 m.
Distribution: U 1, 3; K 1e4; T 2, 7; Eastern Zaire (Kivu). At Kili-
manjaro widespread.
Pollination: birds and insects (Steenhuisen et al., 2012).
5.2.63. Ranunculaceae
Perennial herbs, often with more or less developed rhizome,
sometimes annual or biennial herbs, halfshrubs or lianae. The
family prefers the temperate or cool climate and is rare in the
tropics. The family comprises 59 genera and ca. 2500 species
(Tamura, 1993).
5.2.63.1. Pollination. Specialized and non-specialized insects; most
of the euphilic Ranunculaceae are bee-pollinated, many Delphi-
nieae and Aquilegia spp. typically bumblebee-pollinated. In both
groups also adaptive radiations occurred towards hawkmoth and
bird pollination (Tamura, 1993, and literature herein).
5.2.63.2. Pollen morphology. Pollen grains in Ranunculaceae are 3-
colpate, pantocolpate or pantoporate. The sculpture of the grains is
remarkably uniform (Nowicke and Skvarla, 1979): the only excep-
tions to a (micro) echinate/perforate tectum are found in Hellebo-
rus, Trollius, Hydrastis and Kingdonia (striate tectum) (Nowicke and
Skvarla, 1982). The exine is often underlain by an endexine of
considerable thickness, while the columellae are often large in size,
sparsely distributed, or strongly reduced so that the tectum and the
foot layer are scarcely separated.
Clematis brachiata Thunb (Plate 18, 31e32 & Plate 19, 1). SYN
C. hirsuta Guill. & Perr.
Shape: circular, trilobate ~21e24 mm.
Apertures: 3-colpate, colpi (5/6) with obtuse ends.
Exine: tectate, columellate, microechinate.
Habitus: climbing shrubby plant, more rarely decumbent in
grassland, 1e4 m or taller.
Habitat: forest edges and openings and in wooded grassland;
1000e3000 m. Erica excelsa, Hagenia and Juniperus forests and
clearings, also in Chagga homegardens; 1150e3090 m.
Distribution: U 1e4; K 3e7; T 2, 4, 5, 7, 8; widely spread in most
parts of tropical Africa. At Kilimanjaro widespread.
Pollination: not known. Clematis species haven been observed to
be pollinated by bumble-bees (Dohzono and Suzuki, 2002) and
other insects.
Ranunculus oreophytus Delile (Plate 19, 2e4).
Shape: circular, spheroidal ~43 mm.
Apertures: pericolpate, colpi short and recessed with irregular
margins.
Exine: tectate, columellate, microechinate.
Habitus: perennial herb, usually acaulescent and with much
elongated somewhat thickened roots.
Habitat: rock clefts and short grass near streams, and other wet
and boggy places, in upland moor; 2240e4350 m. At Kilimanjaro in
middle e upper montane riverine forests, muddy streamsides in
the (sub) alpine zone; 2310e3960 m.
Distribution: U 2, 3; K 3, 4; T 2, 3, 6, 7; Ethiopia, southern Sudan,
eastern DR Congo. At Kilimanjaro scattered.
Pollination: not known. Generalistic pollination syndrome with
a great variety of insect pollinators is observed in Ranunculus
(Blackmore et al., 1995; Steinbach and Gottsberger, 1995).
Thalictrum rhynchocarpum Quart. -Dill. & A. Rich (Plate 19, 5e6).
Shape: circular, spheroidal ~19 mm.
Apertures: periporate, few sunken, roundish pori Ø 3.5e5 mm
with irregular margins.
Exine: tectate, scabrate.
Habitus: perennial herb with erect stems, 1e4 m tall.
Habitat: glades and undergrowth of upland rain-forest, upland
evergreen bushland on stream sides and upland grassland;
1620e3150 m. At Kilimanjaro in lower-middle montane Cassi-
pourea and Ocotea forests, in upper montane Podocarpus, Erica
excelsa, Hagenia and Juniperus forests and clearings; 1670e3200 m.
Distribution: U 1e3; K 3e5; T 2, 3, 4, 6, 7; widely distributed in
tropical Africa from Ethiopia to Cameroon and Fernando Po and
south to Natal. At Kilimanjaro widespread.
Pollination: not known; within the genus Thalictrum ane-
mophily and entomophily are known (Kaplan and Mulcahy, 1971).
Self-compatibility is also reported (Melampy and Hayworth, 1980;
Lubbers and Christensen, 1986).
5.2.64. Resedaceae
Annual, perennial or biennial herbs or small shrubs. A family
comprising six genera and 70e75 species, distributed in arid re-
gions of the Mediterranean and eastwards to NW India, in E, W and
S Africa, SW United States and Mexico (Kubitzki, 2003b).
5.2.64.1. Pollination. Insects; some species are also self-fertile
(Kubitzki, 2003b, and literature herein).
5.2.64.2. Pollen morphology. Pollen grains are 3-colpate (-colpor-
idate), prolate spheroidal to prolate. Sexine as thick as nexine or
slightly thicker or thinner, finely reticulate or microperforate
 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386 369

Plate 19. RANUNCULACEAE: 1 Clematis brachiata, 2e4 Ranunculus oreophytus, 5e6 Thalictrum rhynchocarpum, RESEDACEAE: 7e9 Caylusea abyssinica, RHAMNACEAE: 10e12 Scutia
myrtina, ROSACEAE: 13e16 Alchemilla volkensii, 17e18 Hagenia abyssinica, RUBIACEAE: 19e21 Chassalia kenyensis, 22e25 Chassalia parviflora, 26e28 Galiniera saxifraga, 29e31
Kohautia coccinea, 32e34 Lasianthus kilimandscharicus, 35 Mussaenda microdonta.

(Erdtman, 1952; Ramos, 1996). Habitat: weed of secondary grassland, abandoned cultivations,
Caylusea abyssinica (Fresen.) Fisch. & C.A. Mey (Plate 19, 7e9). waste places, riversides, etc.; 1200e3000 m. At Kilimanjaro on
Shape: subprolate ~20  23 mm, trilobate. roadsides, fallow arable land, waste places; 1000e1500 m.
Apertures: 3-colpate, colpi (5/6) tenuimarginate with granulate Distribution: U 1, 2; K 1, 3e6; T 2, 5e7; Eritrea, Ethiopia. At
margins, wider at equator, with acute ends. Kilimanjaro scattered.
Exine: tectate, finely reticulate. Pollination: not known; maybe insects as bees have been
Habitus: erect or ascending herb, occasionally bushy, 1.5e10 m observed on the flowers.
high.
370 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
5.2.65. Rhamnaceae
Deciduous or evergreen, often thorny trees, shrubs, woody
climbers or lianas, rarely herbs. An almost cosmopolitan family of
52 genera and about 925 species (Medan and Schirarend, 2004).
5.2.65.1. Pollination. Generalized, unspecialized entomophily is the
rule, with Hymenoptera and Diptera co-dominating nearly always
(Medan and Schirarend, 2004).
5.2.65.2. Pollen morphology. Pollen grains are usually radial sym-
metrical, angulaperturate, 3 (4)-zono-colporate, suboblate to sub-
prolate and angular to circular in polar view. Apertures are
composed of comparatively narrow colpi and ±narrow, lalongate to
elliptic, rarely lolongate endoapertures. The colpi extend over about
4/5 of the length of the polar axis. The sexine is mostly pertectate-
perforate. The tectum is microreticulate, striate, regulate, fossulate,
verrucate, pilate or ± psilate, with ±densely spaced perforations.
The nexine is always well differentiated, thin at the mesocolpia, and
±thickened at the colpi (Erdtman, 1952; Medan and Schirarend,
2004).
Scutia myrtina (Burm. f.) Kurz (Plate 19, 10e12).
Shape: oblate, triangular ~25 mm.
Apertures: 3-colporate, colpi (4/6e5/6) very narrow, pori rect-
angular broad, pori protruding (especially in polar view) due to
thickened exine (annulus?).
Exine: tectate, reticulate.
Habitus: shrubs or rarely small trees 2e5 (10) m tall, usually
scandent.
Habitat: in a wide variety of situations from forest margins to
bushland, thicket and wooded grassland; from sea-level to 2700 m.
At Kilimanjaro in dry and riverine colline and submontane forests,
lower and middle montane Cassipourea forests; 850e2130 m.
Distribution: U 1e4; K 1e7; T 1e7; Z; DR Congo, Burundi,
Zambia, Zimbabwe, Malawi, Mozambique, South Africa,
Madagascar, Seychelles, Mascarene Is., Ceylon, India, Burma,
Thailand, North Vietnam. At Kilimanjaro widespread in drier forest
types of lower areas.
Pollination: insects (bees, wasps, flies) (Martins, 2014).
5.2.66. Rosaceae
Woody or herbaceous. A moderately large, almost cosmopolitan
family with 85 genera and ca. 2000 sexual species; apart from
those, there are a large number of obligately or facultatively
apomictic ‘microspecies’ in several genera (Kalkman, 2004).
5.2.66.1. Pollination. Pollinating insects are especially flies and
short-tongued bees like Anthrena but, to a lesser degree, also long-
tongued bees, beetles and even butterflies (Kalkman, 2004, and
literature herein).
5.2.66.2. Pollen morphology. Pollen grains do not display large
variation; they are monads, more or less spheroidal, oblate to
prolate, and generally 3-colporate, sometimes 3 colporidate. The
colpi are usually relatively long. In some Rosaceae the ectoapter-
ures are operculate (Erdtman, 1952; Kalkman, 2004).
Alchemilla volkensii Engl (Plate 19, 13e16).
Shape: circular, subprolate, ~20  25 mm.
Apertures: 3-colporate, colpi (4/5) wide with obtuse ends, pori
elliptic tall but inconspicuous.
Exine: tectate, scabrate/microreticulate, slightly striate.
Habitus: herb with central rosette and creeping stolons, at in-
tervals rooting at the nodes and producing tufts of basal leaves and
erect flowering stems.
Habitat: upland rain-forest, moist bamboo-thickets, upland
evergreen bushland, often covering the ground in damp places;
1500e2900 m. At Kilimanjaro mainly in upper montane Podo-
carpus, Erica excelsa, Hagenia and Juniperus forests, also on montane
forest tracks, grasslands and streamsides; 1320e3950 m.
Distribution: U 2, 3; T 2; not known elsewhere. At Kilimanjaro
widespread.
Pollination: probably apomictic and/or pollinated by insects
(Davis et al., 1906).
Hagenia abyssinica J.F. Gmel (Plate 19, 17e18).
Shape: broad rhombic, convex triangular ~32 mm.
Apertures: 3-colporate, lalongate pori with operculum, cuspidal
colpi 1/3 in length.
Exine: tectate, reticulate.
Habitus: rather slender tree up to 20 m tall.
Habitat: upland rain-forest, often above the moist bamboo-
thickets, and in upland evergreen bushland; 2400e3600 m. At
Kilimanjaro mainly in upper montane Podocarpus, Erica excelsa,
Hagenia and Juniperus forests; 1600e3300 m.
Distribution: U 1e3; K 3e5; T 2, 3, 7; DR Congo, Ethiopia, Sudan
and also northern Zimbabwe and Malawi. At Kilimanjaro wide-
spread in the upper forest belt.
Pollination: wind (Feyissa et al., 2007).
5.2.67. Rubiaceae
Small to large trees, shrubs or less often annual or perennial
herbs or woody or herbaceous climbers. A large family of about 500
genera and 6000 species, predominantly tropical and adapted to
moist environments (Verdcourt et al., 1976).
5.2.67.1. Pollination. Most species are insect pollinated, many
having white flowers strongly scented at night, being obviously
pollinated by moths. Anthospermum with its long styles and
dangling anthers is wind pollinated, and a few of the larger flow-
ered Vanguerieae may even be bird pollinated. Doubtless, despite
the special mechanisms, many cases of self-pollination exist
(Verdcourt et al., 1976).
5.2.67.2. Pollen morphology. Pollen grains 3 e polycolpate, 2 e 4-
colporate or 3-porate (exceptionally polycolpate or non-
aperturate), peroblate eprolate, united in tetrads in some species
(Erdtman, 1952).
Chassalia kenyensis Verdc (Plate 19, 19e21).
Shape: circular, spheroidal ~44 mm.
Apertures: 4-colpate, colpi short (1/2) and relatively narrow but
often lacerated, not clearly separable from the reticulum.
Exine: tectate, reticulate, heterobrochate.
Habitus: small erect or somewhat scandent shrub 0.9e1.8 m tall,
with much-branched ridged stems.
Habitat: evergreen forest; 1650e2250 m. At Kilimanjaro in
lower montane Newtonia gorge forests; 1600e1640 m.
Distribution: K 4; T 2, ?3; not known elsewhere. At Kilimanjaro
rare.
Pollination: probably insects like in other Chassalia species (e.g.
Pailler et al., 1998).
Chassalia parviflora Benth (Plate 19, 22e25).
Shape: circular, spheroidal ~46 mm.
Apertures: 4-colpate, colpi very short (1/5) and relatively nar-
row but often lacerated, not clearly separable from the reticulum.
Exine: tectate, reticulate, heterobrochate.
Habitus: much-branched bushy shrub to small tree 2e5.5 (7.5)
m tall.
Habitat: evergreen forest including rain- and mist-forest and
also drier types; 600e2300 m. At Kilimanjaro in lower-middle
montane Cassipourea and Ocotea forests; 1680e2600 m.
 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
Distribution: K 7; T 2, 3, 6e8; N Malawi, NE Zambia. At Kili-
manjaro widespread.
Pollination: probably insects like in other Chassalia species (e.g.
Pailler et al., 1998).
Galiniera saxifraga (Hochst.) Bridson (Plate 19, 26e28).
Shape: circular, spheroidal, ~18 mm.
Apertures: 3-colporate, long (5/6) and narrow colpi, pori rect-
angular to elliptic broad with thick annulus.
Exine: tectate, very thick, stratification difficult to see, scabrate.
Habitus: shrub or small tree 1.8e14 m tall.
Habitat: forest; (760-) 1700e3000 m. At Kilimanjaro mainly in
lower-middle montane riverine, Cassipourea and Ocotea forests;
1560e2900 m.
Distribution: U 1e4; K 3e6; T 2e4, 6, 7; Zaire, Rwanda, Burundi,
Sudan, Ethiopia, Malawi, Zambia. At Kilimanjaro widespread.
Pollination: probably insects, but no further information
available.
Kohautia coccinea Royle (Plate 19, 29e31).
Shape: circular, spheroidal ~21 mm.
Apertures: 6-colpate, very narrow colpi (3/5 to 4/5) with acute
ends.
Exine: tectate, coarsely reticulate, heterobrochate.
Habit: annual erect, unbranched or sparsely branched herb
0.07e0.45 (0.7) m tall.
Habitat: grassland, including seasonally wet areas, grassland
with scattered trees, woodland edges, gravel paths, waste ground,
shallow soil on rock outcrops and also as a weed in cultivated fields,
by roadsides, etc.; 880e2400 m. At Kilimanjaro in dry savanna
grasslands; 1220 m.
Distribution: U 1e4; K 1e6; T 1, 2, 4, 7, 8; Z; Senegal, N. Nigeria,
Zaire, Ethiopia, Malawi, Zambia, Zimbabwe, Mozambique, also in N.
India. At Kilimanjaro rare.
Pollination: Insects. Pollination mechanism in the genuas
Kohautia conspicuously specialized, or unspecialized (Groeninckx
et al., 2010).
Lasianthus kilimandscharicus K. Schum (Plate 19, 32e34).
Shape: circular to subtriangular ~35 mm, subprolate.
Apertures: 3-porate, pori rectangular and large ~3  7 mm.
Exine: distinctly tectate, (meso) reticulate.
Habit: Shrub or small tree 1.2e7.5 m tall.
Habitat: Understorey of upland rain-forest, often dominant
particularly in Ocotea forests; 1710e2400 m. At Kilimanjaro in
lower-middle montane Cassipourea and Ocotea forests;
1590e2650 m.
Distribution: U 2; K 1, 3e5, 7; T 2, 3, 5e7; Zaire, Burundi,
Mozambique, Malawi, Zambia, Zimbabwe. At Kilimanjaro
widespread.
Pollination: Insects, no further details known.
Mussaenda microdonta Wernham (Plate 19, 35 & Plate 20, 1).
Shape: circular, spheroidal ~22 mm.
Apertures: 5-porate, pori Ø ~5 mm ± circular with annulus
(endocosta).
Exine: tectate, reticulate, heterobrochate.
Habitus: shrub or small tree 3e9 (-? 27) m tall.
Habitat: evergreen forest; (1000-)1120e2130 m. At Kilimanjaro
in lower montane Hallea riverine and Newtonia gorge forests;
1400e1930 m.
Distribution: U 3; K 4, 5; T 2; not known elsewhere. At Kili-
manjaro rare.
Pollination: not known for this species. Other Mussaenda species
are pollinated by butterflies, birds, bees, hawkmoths (e.g. Borges
et al., 2003; Chen et al., 2013).
Pavetta abyssinica Fresen var. abyssinica (Plate 20, 2e4).
Shape: circular, spheroidal/subprolate ~25  29 mm.
Apertures: 3-colporate, long straight colpi (5/6) with obtuse
371
ends, slightly constricted at equator, pori large, rhombic broad with
annulus.
Exine: tectate, reticulate.
Habitus: Bush or small tree, sometimes climbing, 1e7 (9) m
tall.
Habitat: Forest or occasionally scrub; 1050e2500 m. At Kili-
manjaro in lower-middle montane Cassipourea and Ocotea forests;
1580e2400 m.
Distribution: U 1, 3; K 1e6; T 2, 3, 5e7; ?Sudan, Ethiopia. At
Kilimanjaro widespread.
Pollination: not known for this species; other Pavetta species are
known to be pollinated by butterflies (Balasubramanian, 1993).
Pentas lanceolata (Forssk.) Deflers subsp. lanceolata var. lanceo-
lata (Plate 20, 5e9).
Shape: subprolate/rectangular tall ~21  28 mm, circular.
Apertures: 3-colporate, narrow colpi (4/6); large, rectangular
broad pori, probably zonorate.
Exine: tectate, finely reticulate.
Habitus: herb or subshrub with erect or straggling mostly
woody stems 0.5e1.3 m tall.
Habitat: grassland, bushland, thicket, also in evergreen forest,
sometimes on rocky cliffs and escarpments; 1440e3000 m. At
Kilimanjaro in colline e middle montane riverine forests, upper
montane Podocarpus and Juniperus forests, also in open habitats,
e.g. dry submontane Hyparrhenia grasslands; 970e2900 m.
Distribution: U 1; K 1e4, 6, T 2; also in Arabia, Ethiopia, Sudan
and Afars Issas. At Kilimanjaro widespread.
Pollination: pollinated by a small nitidulid beetle of the genus
Meligethes (Bahadur, 1970).
Pentodon pentandrus (Schumach. & Thonn.) Vatke var. pen-
tandrus (Plate 20, 10e12).
Shape: circular, spheroidal ~25 mm.
Apertures: 3-colporate, colpi (3/5) wider at equator and with
acute ends, broad rectangular pori.
Exine: tectate, reticulate.
Habitus: usually annual or short-lived perennial herb, with
weak decumbent, procumbent or rarely suberect, often single
stems 0.04e0.9 m long.
Habitat: Swamp, lake and river margins, seasonally wet ground,
muddy hollows, etc.; (? 200-) 450e2250 m. At Kilimanjaro in col-
line swamps on lake shores; 800 m.
Distribution: U 2e4; K 1, ?4, ?5, ?7; T 1e4, 6, 7, 8; widespread in
tropical Africa from Cape Verde Is., W. Africa (Senegal e Angola),
Arabia, Sudan, Ethiopia and Somali Republic to Malawi and Zambia,
Zimbabwe, Botswana, South West Africa, Madagascar, also in USA
(Florida and Texas), Cuba, Nicaragua and Brazil. At Kilimanjaro rare.
Pollination: not known.
Psychotria capensis subsp. riparia (K. Schum. & K. Krause) Verdc
(Plate 20, 13e14). SYN P. riparia.
Shape: circular, spheroidal ~34 mm (often folded).
Apertures: 3-colporate, colpi short (1/5) and inconspicuous, pori
small.
Exine: tectate, ectexine thicker than endexine, coarsely reticu-
late, heterobrochate.
Habitus: shrub 2.5e4.5 m tall, or tree up to 6 (20) m.
Habitat: evergreen bushland, coastal bushland, rocky places,
wooded grassland and forest-edges and especially riparian forest
(often with Phoenix). At Kilimanjaro mainly in dry savanna forests
and colline e submontane riverine forests; 780e1370 m.
Distribution: U 1, 3; K 4, 7; T ?1, 2e8; Z; P; Mozambique, Zambia
and Malawi. At Kilimanjaro scattered.
Pollination: not known; Psychotria species are generally polli-
nated by bees, flies, moths and birds (Castro and Araujo, 2004;
Virillo et al., 2007).
Psychotria cyathicalyx E.M.A. Petit (Plate 20, 15e16).
372 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386

Plate 20. RUBIACEAE: 1 Mussaenda microdonta, 2e4 Pavetta abyssinica var. abyssinica, 5e9 Pentas lanceolata ssp. lanceolata var. lanceolata, 10e12 Pentodon pentandrus var. pen-
tandrus, 13e14 Psychotria capensis ssp. riparia, 15e16 Psychotria cyathicalyx, 17e18 Psychotria fractinervata, 19e20 Psychotria petiginosa, 21e22 Richardia scabra, RUTACEAE: 23e25
Calodendrum capense, 26e28 Clausena anisata, 29e31 Toddalia asiatica, SANTALACEAE: 32e35 Osyris lanceolata, SAPINDACEAE: 36e37 Allophylus ferrugineus, 38e39 Allophylus
rubifolius var. rubifolius.

Shape: circular/trilobate, prolate ~18  26 mm.
Apertures: 3-colporate, long colpi (5/6) narrow, pori rectangular
broad.
Exine: tectate, coarsely reticulate.
Habit: Shrub 2e5 m tall or tree up to 10 m tall.
Habitat: upland evergreen forest, including Ocotea forest and
Podocarpus-Erica associations at upper edges, often dominant
where it occurs; 1600e3000 m. At Kilimanjaro mainly in middle
montane Ocotea forests and upper montane Podocarpus forests;
1880e2920 m.
Distribution: T 2, 3, 6; not known elsewhere. At Kilimanjaro
widespread.
Pollination: not known; Psychotria species are generally polli-
nated by bees, flies, moths and birds (Castro and Araujo, 2004;
Virillo et al., 2007).
Psychotria fractinervata E.M.A. Petit (Plate 20, 17e18).
Shape: convex triangular, spheroidal ~53 mm.
Apertures: 3-colporate, colpi (2/5) broad, colpi margins not
smooth, pori large and unclear in shape, wide open in polar view.
Exine: tectate, thick exine, coarsely reticulate, heterobrochate.
Habitus: shrub or small tree 2e6 (7.5) m tall.
Habitat: upland rain-forest, including Podocarpus, bamboo, etc.
and reaching Hypericum zone; (1500-) 1800e2600 m. At Kili-
manjaro mainly in lower e middle montane Ocotea forests;
1590e2710 m.
Distribution: U 3; K 3, 4, 6; T 2; not known elsewhere. At Kili-
manjaro widespread.
Pollination: not known; Psychotria species are generally polli-
nated by bees, flies, moths and birds (Castro and Araujo, 2004;
Virillo et al., 2007).
Psychotria petiginosa Brenan (Plate 20, 19e20).
Shape: circular, spheroidal ~70 mm.
Apertures: 3-colpate, short (2/6) colpi wide with acute ends,
margins not clearly distinguishable.
Exine: tectate, coarsely reticulate, ±homobrochate.
Habitus: shrub.
 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
Habitat: floor of evergreen forest; 1500e2000 m. At Kilimanjaro
mainly in lower e middle montane Cassipourea forests and lower
montane Ocotea forests; 1710e2410 m.
Distribution: T 2, 3; not known elsewhere. At Kilimanjaro
widespread.
Pollination: not known; Psychotria species are generally polli-
nated by bees, flies, moths and birds (Castro and Araujo, 2004;
Virillo et al., 2007).
Richardia scabra L (Plate 20, 21e22).
Shape: circular, spheroidal ~86 mm.
Apertures: stephanocol(por)ate, 21 very short colpi, presence of
pori not clear.
Exine: tectate, bacculate.
Habitus: perennial (or? annual) prostrate herb, often forming a
mat; stems 0.05e0.55 m long.
Habitat: cultivations, newly cleared ground, roadsides on red
soil in forest clearings, sandy grassland and bushland; 0e1600 m.
At Kilimanjaro in colline savanna e lower montane grasslands, also
in disturbed places (Chagga homegardens, roadsides, fallow arable
fields and waste places); 820e1900 m.
Distribution: T2e4, 6; Zimbabwe, South Africa, S. and Central
America, Jamaica, Cuba and USA. At Kilimanjaro widespread.
Pollination: bees have been observed on this species (unpub-
lished). Other Richardia species are pollinated by various insects
such as Coleoptera, Diptera, Hymenoptera, and Lepidoptera (Cruz
and Martins, 2015).
5.2.68. Rutaceae
Trees or shrubs, sometimes scandent, rarely herbs. A family of
154 genera and about 2100 species; nearly cosmopolitan, but
mainly tropical and subtropical; most divers in Australasia
(Kubitzki et al., 2011).
5.2.68.1. Pollination. Entomogamous, with smaller Hymenoptera
and Diptera as predominating pollinators in the humid forest
biome. Calodendrum is pollinated by butterflies and bees, bees also
take part in the pollination of Toddalia (Kubitzki et al., 2011).
5.2.68.2. Pollen morphology. Pollen grains are usually 3-colporate
and prolate with a reticulate tectum and lalongate endoapertures.
This is the widespread condition, which usually is only moderately
modified: in Rutoideae, exine sculpture relatively often tends to be
striate, less often microperforate or very coarsely reticulate, rarely
echinate or baculate (Erythrochiton, some Angostura, Nycticalanthus,
Spiranthera), and pollen in the Angostura Alliance is sometimes
spherical and then often has 4e6, very short colpi (some Angostura,
Galipea, Sigmatanthus, Ticorea) (Grant et al., 2000; Kubitzki et al.,
2011).
Calodendrum capense (L.f.) Thunb (Plate 20, 23e25).
Shape: circular, spheroidal ~24 mm.
Aperture: 3-porate, pori circular with annulus slightly
protruding.
Exine: tectate, microreticulate.
Habitus: tree up to 20 m high.
Habitat: upland evergreen and riverine forest; 1200e2200 m. At
Kilimanjaro in submontane Croton-Calodendrum forests;
1300e2030 m.
Distribution: U 2, 4: K 1, 3, 4, 6, 7; T 2, 5, 6; Malawi, Zimbabwe,
South Africa. At Kilimanjaro on the western and northern slope
widespread.
Pollination: butterflies and bees (Kubitzki et al., 2011).
Clausena anisata (Willd.) Benth (Plate 20, 26e28).
Shape: subprolate/rhombic tall ~21  29 mm, subtriangular.
Aperture: 3-colporate, lalongate pori narrow (rectangular),
straight colpi 5/6 with margo.
373
Exine: tectate, striato-reticulate, slightly thickened around the
pore-colpus transition.
Habitus: shrub or small tree up to 4 (10) m high.
Habitat: forests, often margins and regeneration, persisting in
bushland and wooded grassland; 1e2450 (2700) m. At Kili-
manjaro in colline riverine forests, submontane Croton-Caloden-
drum forests, lower e middle montane Cassipourea forests and
upper montane Juniperus forests; 900e2990 m.
Distribution: U 1e4; K 1, 3e7; T 1e8; Z; P; Guine e to Ethiopia
and south to Cape Province of South Africa. At Kilimanjaro wide-
spread in drier forest types.
Pollination: insects (Kubitzki et al., 2011).
Toddalia asiatica (L.) Lam (Plate 20, 29e31).
Shape: subprolate ~20  24 mm, trilobate.
Aperture: 3-colporate, pori lalongate (nearly zonoporate), colpi
(4/5) narrow, slightly wider at equator, with acute ends.
Exine: tectate, reticulate, high reticulum with relatively large
lumina, ± homobrochate.
Habitus: scrambling or climbing shrub.
Habitat: forest edges, bushland and wooded grassland;
1e2800 m. At Kilimanjaro in colline riverine forests, submontane
Croton-Calodendrum forests, lower e middle montane Cassipourea
forests and upper montane Juniperus forests, also in Chagga
homegardens; 780e2450 m.
Distribution: U 1e4; K 1e7; T 1e8; Z; Zaire, Rwanda, Sudan,
Ethiopia, Malawi, Mozambique, Zimbabwe, Zambia, South Africa,
Madagascar, Mascarene Is., India. At Kilimanjaro widespread in
drier forest types.
Pollination: bees and other insects (Kubitzki et al., 2011).
5.2.69. Santalaceae
Trees, shrubs or herbs, all assumed or known to be semi-
parasitic. A family of about 36 genera and several hundred species,
throughout the temperate and tropical regions (Kuijt et al., 2015).
5.2.69.1. Pollination. Probably bees, flies and beetles (Kuijt et al.,
2015).
5.2.69.2. Pollen morphology. Pollen grains are subisopolar or het-
eropolar, 3 aperturate, oblate to prolate (Erdtman, 1952). Sculp-
turing is highly variable, ranging from psilate through tuberculate
and areolate to spinulate, echinate-perforate as well as murciate-
striate (Lobreau-Callen, 1982).
Osyris lanceolata Hochst. & Steud (Plate 20, 32e35).
Shape: triangular/trilobate, spheroidal ~19 mm.
Apertures: 3-colporate, pori elliptic/rectangular broad, colpi (4/
5) narrow and slightly constricted at equator.
Exine: tectate, scabrate.
Habitus: shrub or small tree, 1.5e9 (14) m tall.
Habitat: upland dry evergreen forest and mist forest, with
associated bushland and grassland, extending down rivers and
from there marginally into deciduous woodland; (50-)
900e2700 m. At Kilimanjaro in dry savanna forests and woodlands
and thicked edges in dry submontane Hyparrhenia grasslands;
1000e1730 m.
Distribution: U 1e3; K 1e7; T 1e8; widespread in Africa from
Algeria to Ethiopia and South Africa, Europe (Iberian Peninsula,
Balearic Is.), Asia (India to China), Socotra. At Kilimanjaro
widespread.
Pollination: probably insects like in other Osyris species with
small green flowers (Aronne et al., 1993).
5.2.70. Sapindaceae
Trees, treelets, shrubs, lianas or herbaceous climbers. A mostly
tropical to subtropical family, with a few genera extending to sub-
374 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
temperate zones; 141 genera and about 1900 species (Acevedo-
Rodríguez et al., 2011).
5.2.70.1. Pollination. Primarily bee-pollinated, flowers can also be
attractive to other kinds of pollinators including Lepidoptera and
humming birds (Acevedo-Rodríguez et al., 2011).
5.2.70.2. Pollen morphology. Pollen grains are usually isopolar or
subisopolar monads. Grain size is usually between 20 and 30 mm,
and grains are oblate to prolate in shape. Colporate pollen is usually
suboblate to prolate, whereas pollen with small apertures (porate,
brevicolporate) or with connected apertures (syncolporate, para-
syncolporate) has a more oblate shape. The equatorial outline is
almost circular to bluntly triangular. Generally, pollen grains are 3-
colporate, but often small percentages of 2 colporate and 3e4-
porate grains co-occur (Erdtman, 1952; Acevedo-Rodríguez et al.,
2011 and references herein).
Allophylus ferrugineus Taub (Plate 20, 36e37).
Shape: triangular (quadrangular when 4-porate) ~34 mm, oblate.
Apertures: 3 (4) porate, exine thicker around pori, exine below
pori bent inwards, vestibulum.
Exine: tectate, scabrate.
Habitus: tree or shrub or sometimes a climber or creeper
0.9e9 m long or tall.
Habitat: forest, often by streamsides, hillside thicket, sometimes
on termite mounds, plantations; (650-) 1050e2400 (-?2700) m. At
Kilimanjaro mainly in lower montane Cassipourea and Ocotea for-
ests; 960e2650 m.
Distribution: U 2e4; K 3e6; T 1, 2, 4, 6, 7; Cameroon, Central
African Republic, Zaire, Rwanda, Sudan, Ethiopia and Angola. At
Kilimanjaro widespread.
Pollination: probably bee pollinated like in other Allophylus
species (Aluri et al., 1998).
Allophylus rubifolius (Hochst. ex A. Rich.) Engl. var. rubifolius
(Plate 20, 38e39).
Shape: triangular ~21 mm, (sub) oblate.
Apertures: 3-porate, pori circular with vestibulum, exine around
pori not thickened.
Exine: tectate, faintly striato-microreticulate.
Habitus: shrub or small to medium-sized tree 3e7 (12) m tall,
also occasionally described as a creeper.
Habitat: grassland with scattered trees, rough grassland, thicket,
edges of cultivations, woodland, sometimes riverine; ?0e2250 m.
At Kilimanjaro in dry woodlands and riverine forests in the colline
savanna zone and submontane Croton-Calodendrum forests;
780e2070 m.
Distribution: U 1e4; K 1e7; T 1e8; E Zaire, Sudan, Ethiopia and
N Somalia, south to South Africa. At Kilimanjaro scattered.
Pollination: probably bee pollinated like in other Allophylus
species (Aluri et al., 1998).
Filicium decipiens (Wight & Arn.) Thwaite (Plate 21, 1e3).
Shape: subtriangular, circular ~ 21 mm.
Aperture: 3-colporate, pori elliptic/rectangular broad with ves-
tibulum, exine thicker around pore and straight or bent inwards
below the pore, colpi (4/5) narrow.
Exine: tectate, bacculate.
Habitus: slender tree 4.5e20 (30) m tall.
Habitat: mixed forest in river valleys; 450e1600 m. At Kili-
manjaro in colline e submontane riverine and gorge forests;
0.960e1450 m.
Distribution: K 1, 4, 7; T 2, 3, 6, 7; Ethiopia, Malawi,
Mozambique, Zimbabwe, Madagascar, Comoro Is., India and Sri
Lanka, also cultivated in Nairobi (arboretum) & Amani (Boma
Garden), Java, Sabah, Fiji, Samoa and Hawaii. At Kilimanjaro rare.
Pollination: not known; most likely insects as other
Sapindaceae.
Paullinia pinnata L (Plate 21, 4e5).
Shape: triangular ~ 44 mm, (sub) oblate, but large variation in
size.
Aperture: 3 (4) porate, large elliptic pori like a flattened cone Ø
~4 mm.
Exine: tectate, finely reticulate, seemingly homobrochate,
endexine encloses the pori more than the ectexine.
Habitus: shrubby climber 2.5e8 (-? 15) m high.
Habitat: forest margins, gallery forest, moist thicket and scrub;
0e1600 m. At Kilimanjaro in colline e submontane riverine and
gorge forests, aslo in Chagga homegardens; 780e1750 m.
Distribution: U 2e4; K 3e7; T 1e4, 6e8; Z; P; troughout tropical
Africa, Madagascar, tropical and subtropical America. At Kili-
manjaro scattered.
Pollination: bees and wasps (Willemstein, 1987).
5.2.71. Sapotaceae
Trees or shrubs, rarely geoxylic suffrutices or lianas, sometimes
spiny. A pantropical family of 53 genera and about 1100 species,
mostly in humid forest, but some genera (e.g. Sideroxylon, Argania)
extending into semi-arid and arid regions (Pennington, 2004).
5.2.71.1. Pollination. Although the majority of Sapotaceae certainly
is entomophilous, in the literature only instances of bat pollination
seem to have been recorded (Pennington, 2004).
5.2.71.2. Pollen morphology. A stenopalynous family. Pollen grains
3e4 (-5e6) colporate. The colpi are usually narrow, occasionally
broad, and range from vestigial to ca. 4/5 or 5/6 of the polar length.
The pori are narrowly or broadly lalongate, less frequently circular
or, extremely rare, broadly lolongate. The grains are predominantly
prolate-spheroidal, subprolate or prolate, occasionally spheroidal
or, rarely, oblate-spheroidal; the amb is more or less circular. The
endexine is usually absent in the polar region but greatly thickened,
especially in the area surrounding the endoaperture and underly-
ing the colpus (Erdtman, 1952; Pennington, 2004).
Aningeria adolfi-friedericii (Engl.) Robyns & G.C.C. Gilbert agg
(Plate 21, 6e8).
Shape: subprolate-rectangular, circular, ~27  36 mm.
Apertures: 3- or 4-colporate, colpi long (5/6) and very narrow;
pori circular, protruding due to thickened exine.
Exine: tectate, exine thicker at equator and around apertures,
psilate.
Habitus: tall tree up to 50 m high. Long straight ± fluted bole and
buttressed base.
Habitat: upland rain-forest, frequently associated with Podo-
carpus, rarely in riverine forest; 1430e2500 (?3200) m. At Kili-
manjaro in lower montane Cassipourea and Newtonia gorge forests;
1650e2080 m.
Distribution: T 2, 3, 4, 7, 8; K 3, 4, 5; U 1e3; extreme east of DR
Congo, Rwanda and SW Ethiopia, ?Sudan. At Kilimanjaro rare.
Pollination: probably bees and other insects like in other Anin-
geria species (e.g. Roubik and Villanueva-Gutie rrez, 2009;
Muradian and Rival, 2013).
5.2.72. Solanaceae
Annual or perennial erect or climbing herbs, shrubs or small
trees. About 96 genera with some 2300 species, cosmopolitan with
exception of polar regions. Greatest species concentration and di-
versity occurs in Central and South America. Within Africa, the
family is relatively poorly represented. Nevertheless, it is probable
that tropical East Africa contains the largest number of solanaceous
species found in Africa (Edmonds, 2012).
 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386 375

Plate 21. SAPINDACEAE: 1e3 Filicium decipiens, 4e5 Paullinia pinnata, SAPOTACEAE: 6e8 Aningeria adolfi-friedericii agg., SOLANACEAE: 9e11 Solanum nigrum, 12e14 Solanum
seaforthianum, STERCULIACEAE (Malvaceae): 15e16 Dombeya burgessiae, 17e19 Leptonychia usambarensis, THYMELIACEAE: 20e21 Gnidia apiculata, 22e23 Gnidia glauca, 24e25
Gnidia subcordata, TILIACEAE (Malvaceae): 26e28 Grewia bicolor, 29e31 Grewia microcarpa, 32e34 Triumfetta brachyceras, 35e37 Triumfetta flavescens, ULMACEAE (Cannabaceae):
38e40 Celtis gomphophylla.

5.2.72.1. Pollination. Pollinators range from bees, moths, insects in
general to birds and bats (Knapp, 2010).
5.2.72.2. Pollen morphology. Pollen grains (2-) 3e5 (6) colpate,
-colporidate, (zono)-colporate (or colporidate), sometimes ina-
perturate, oblate - prolate (Erdtman, 1952). Exine sculpture can be
rugulate, scabrate, echinate and reticulate and the stratification
consists of sexine and nexine (Persson et al., 1994).
Solanum nigrum L (Plate 21, 9e11).
Shape: subtriangular, spheroidal to suboblate, ~20 mm.
Apertures: 3-colporate, colpi straight (5/6) with fissured mar-
gins, pori lalongate with vestibulum, broader than colpi, exine not
thickened around pori.
Exine: tectate, psilate.
Habitus: annual to perennial herb, erect, decumbent or pros-
trate, 0.1e1.1 m high. Main stems sometimes woody, spreading up
to 2 m.
Habitat: old or secondary cultivation areas, grassland, forest
edges and in clearings or on stream banks in upland forests, on
mountain tops and on rocky hillsides with Brachystegia, Acacia and
Commiphora; (15-)1200e2200 m. At Kilimanjaro mainly in Chagga
homegardens, used as vegetable; 960e2310 m.
Distribution: U 1e3; K 3, 4, 6; T 1e3, 5e8; Sierra Leone, Ghana,
Nigeria, Cameroon, Bioko, Sudan, Eritrea, Ethiopia, Somalia, Angola,
Zambia, Malawi, Mozambique, Zimbabwe, Namibia, Botswana,
south Africa, Madagascar, Madeira, Azores, Europe, Middle East,
Asia, introduced and often naturalized in North America, Australia
and New Zealand. At Kilimanjaro widespread.
Pollination: usually self-pollinating but outcrossing can occur
(e.g. Venkateswarlu and Rao, 1972).
Solanum seaforthianum Andrew (Plate 21, 12e14).
Shape: subtriangular, spheroidal ~15 mm.
Apertures: 3-colporate, syncolpate, pori lalongate with oper-
culum, pori slightly protruding, exine thickened around pori, colpi
narrowing towards poles.
Exine: tectate, psilate.
376 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
Habitus: vine, creeper, or trailing to scrambling shrub reaching
7 m high. Climbing by means of twining petioles. Stems woody
basally.
Habitat: semi-deciduous forest, groundwater forest, grassland,
farmland, plantations, forest reserves and gardens; 750e1550 m. At
Kilimanjaro in colline riverine forests and Chagga homegardens;
900e1340 m.
Distribution: U 1; K 4, 7; T 2, 3, 6; native of the West Indies and C
America, now widely cultivated and a common and often natural-
ized escape in Senegal, Sierra Leone, Ghana, Nigeria, DR Congo-
Kinshasa, Zambia, Malawi, Zimbabwe, Namibia, Botswana, South
Africa, Egypt, Madagascar, Mauritius, Reunion, Comoro Islands,
Australia, New Guinea. At Kilimanjaro naturalized, scattered.
Pollination: Pollination in Solanum flowers is performed by in-
sects (mainly bees). The species S. seaforthianum is reported d by
Wagner et al. (1999) as self-compatible.
5.2.73. Sterculiaceae (today Malcaveae Sterculiodeae)
Species described here under former family of Sterculiaceae
today belong to the family of Malvaceae. Therefore, detailed in-
formation is given for Malvaceae or the correspondant subfamily
(see also Malvaceae section of this atlas).
Members of the subfamily of Dombeyoideae (Dombeya) are
mostly shrubs, more rarely herbs or small trees, some of which
grow in more or less xeric habitats. All species are exclusively
palaeotropical. Species of Byttnerioideae (Leptonychia) have a
pantropical distribution; some genera occur in rain-forests (Bayer
and Kubitzki, 2003; Stevens, 2015).
5.2.73.1. Pollination. Species with small flowers within the
Byttnerioideae are pollinated by a variety of insects, whereas the
species with larger flowers are commonly pollinated by vertebrates
e.g. bat or moths. Some Dombeyoideae exhibit secondary pollen
presentation, however pollinators have never been reported
(except for Dombeya, see below) (Bayer and Kubitzki, 2003, and
literature herein).
5.2.73.2. Pollen morphology. Prolate to perprolate, usually reticu-
late grains represent the Grewia-type, which is characteristic of
most Byttnerioideae (except e.g. Leptonychia) and most Grewioi-
deae, both Malvaceae. The Bombax-type (most Bombacoideae and
Leptonychia) has oblate pollen that is angular in polar view and may
be plan- or sinaperturate. Within Bombacoideae, the enormous
variation of shapes, apertures, exine structure and ornamentation
has been described by Nilsson and Robyns (1986). The Malva-type
is globose or rarely (typical for most Malvoideae and Dombeyoi-
deae and some Bombacoideae (Radyera)) somewhat flattened and
covered with ±prominent spines, these may all be of the same or of
different length. Apertures vary from zono-3-colporate to zono-
triporate to zono-spiraporate to pantoporate with up to 300 pori
(all Bayer and Kubitzki, 2003, and literature herein).
Dombeya burgessiae Gerrard ex Harv. & Sond (Plate 21, 15e16).
(Malvaceae Dombeyoideae).
Malva-type.
Shape: circular, subprolate ~45  57 mm.
Apertures: 3-porate, pori circular Ø 4e5 mm with broad annulus
~4e5 mm.
Exine: tectate, echinate, echinae short with acute tip, base of
echinae distinctly broader than tip.
Habitus: small tree or shrub 2e4 (8) m tall.
Habitat: grassland, wooded grassland, riverine thicket and
scrub, open forest and forest edges; (900-) 1500e2000 (2400) m.
At Kilimanjaro in submontane riverine and Croton-Calodendrum
forests; 980e1980 m.
Distribution: U 1e4; K 2e6; T 1, 2, 4, 5, 7, 8; DR Congo-Kinshasa,
Rwanda, Angola, Zambia, Malawi, Mozambique, Zimbabwe,
Swaziland, South Africa. At Kilimanjaro widespread on the western
and northern slope.
Pollination: according to Yeo (1993) the flowers are
melittophilous.
Leptonychia usambarensis K. Schum (Plate 21, 17e19). (Malva-
ceae Byttnerioideae).
Bombax-type.
Shape: triangular (convex) ~31 mm, (sub)oblate.
Apertures: 3-porate, planaperturate, pori elliptic tall, exine
thickened around pori (annulus?).
Exine: tectate, reticulate with large lumina, heterobrochate.
Habitus: shrub or tree, 3e20 m tall.
Habitat: moist forest; 200e350(-1900) m. At Kilimanjaro in
lower montane Newtonia gorge forests; 1350e1780 m.
Distribution: K 4, 7; T 3, 6, 7; Mozambique, Malawi (Misuku
Hills). At Kilimanjaro rare, only on the southern slope.
Pollination: not reported for this species.
5.2.74. Thymeliaceae
Trees, shrubs, lianas, or rarely herbs, evergreen or dedicuous. A
cosmopolitan family of 45 genera comprising about 800 species,
most species and genera occur at latitudes south of 40
N (Herber,
2003).
5.2.74.1. Pollination. Insect pollination seems to predominate
within the family (Herber, 2003), nevertheless, observations of
pollination are rare.
5.2.74.2. Pollen morphology. A stenopalynous family. Pollen grains
usually oligo-polyporate, 20e75 mm in diameter. Sexine is thicker
than nexine, usually provided with vestigial spinules and
patterns ± similar to the “croton-pattern” of Euphorbiaceae
(Erdtman, 1952).
Gnidia apiculata (Oliv.) Gilg (Plate 21, 20e21).
Shape: circular, spheroidal ~21 mm.
Apertures: periporate, circular pori small max. Ø 1.5 mm.
Exine: tectate, very thick exine, scabrate/microreticulate.
Habitus: erect, much branched shrub or undershrub up to 1 m
high.
Habitat: grasslands, dry hillsides, on red soil; 150e1700 m. At
Kilimanjaro in dry colline savanna and submontane Hyparrhenia
grasslands; 1150e1430 m.
Distribution: U 1; K 4, 5; T 2, 3, 5e7; Cameroon, Gabon, Zaire,
Chad, Central African Republic, Sudan. At Kilimanjaro widespread.
Pollination: not known for this species; most Gnidia species are
however pollinated by moths (Manning and Paterson-Jones, 2007).
Gnidia glauca (Fresen.) Gilg (Plate 21, 22e23).
Shape: circular, spheroidal ~34 mm.
Apertures: periporate, circular pori small Ø ~2 mm (slightly
larger than or nearly the size of the lumina).
Exine: tectate, very thick exine, reticulate, heterobrochate.
Habit: large, much-branched shrub up to 3.5 m high or small
tree up to 15 (24) m high.
Habitat: upland forest margins and associated bushland or
wooded grassland; 1500e3300 m. At Kilimanjaro in subalpine
ericaceous bush; 2450e2850 m.
Distribution: U 1, 3; K 1e6; T 2, 4e8; Nigeria, Cameroon, Zaire,
Sudan, Ethiopia, Malawi, Zambia. At Kilimanjaro scattered, only on
the northern slope.
Pollination: not known for this species; most Gnidia species are
however pollinated by moths (Manning and Paterson-Jones, 2007).
Gnidia subcordata Meisn (Plate 21, 24e25).
Shape: circular, spheroidal ~31 mm.
Apertures: periporate, circular pori small Ø ~2 mm, distinctly
 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
larger (about double) than lumina.
Exine: tectate, very thick exine, reticulate, heterobrochate.
Habit: much-branched shrub up to 3.5 m high.
Habitat: upland dry evergreen forest and associated bushland,
Acacia woodland and wooded grassland; 1400e2400 m. At Kili-
manjaro mainly in submontane Croton-Calodendrum forests, also in
dry savanna grasslands; 1150e1850 m.
Distribution: U 1; K 2e7; T 2, 3, 5; Sudan, South Africa. At Kili-
manjaro scattered.
Pollination: not known for this species; most Gnidia species are
however pollinated by moths (Manning and Paterson-Jones, 2007).
5.2.75. Tiliaceae (today Malvaceae Tilioideae).
Species described here under the former family of Tiliaceae
today belong to the family of Malvaceae (here all subfamily Gre-
wioideae). Therefore, detailed information is given for Grewioideae
(see also Malvaceae section of this atlas). Species of Grewioideae
have a pantropical and partly warm temperate distribution; some
genera comprise medium sized trees of humid forests in the Neo-
and/or Paleotropics. Grewia prefers subarid habitats in the Old
World. Corchorus and Triumfetta are represented in all tropical re-
gions with weedy shrubs, halfshrubs or herbs occupying woodland,
savannah or ruderal vegetation (Bayer and Kubitzki, 2003; Stevens,
2015).
5.2.75.1. Pollination. In Grewioideae relatively small flowered
genera point to insect pollination. Larger flowers within this sub-
family are pollinated by vertebrates (Bayer and Kubitzki, 2003).
5.2.75.2. Pollen morphology. Roughly, the pollen types of the Mal-
vaceae can divided into a few main types (Bayer and Kubitzki,
2003): the Grewia-type, the Tilia-type, the Helicteres-type, the
Bombax-type and the Malva-type. The Grewia-type which is mainly
found within the Grewioideae and Byttneroideae prolate to per-
prolate, reticulate grains are representative. Sterculoideae have
less pronounced elongate to spheroidal, mostly coarsely reticulate
pollen grains.
Grewia bicolor Juss (Plate 21, 26e28). (Malvaceae Grewioideae).
Grewia-type.
Shape: prolate ~36  52 mm, circular.
Apertures: 3-colporate, pori rectangular broad ~3  14 mm, colpi
long (4/5), open with obtuse ends.
Exine: tectate, reticulate, heterobrochate.
Habit: shrub or small tree to 6 m tall.
Habitat: woodland, thicket, Commiphora-Acacia bushland,
scattered bush grassland, on dry sandy soil and along streams;
650e1650 m. At Kilimanjaro mainly in dry colline savanna forests
and woodlands; 920e1580 m.
Distribution: U 1; T 1, 2, 4, 5, 7; widespread from Senegal to
Ethiopia, south Botswana, Namibia and NE South Africa. At Kili-
manjaro widespread.
Pollination: insects, mainly by Coleoptera, but also Lepidoptera
and Hymenoptera (Mawdsley and Sithole, 2010).
Grewia microcarpa K. Schum (Plate 21, 29e31). (Malvaceae
Grewioideae).
Grewia-type.
Shape: subprolate ~40  51 mm, circular (often folded) ~50 mm.
Apertures: 3-colporate, colpi (4/5) narrow without distinct
margin, rectangular broad to narrow lalongate pori (sometimes not
very well visible).
Exine: tectate, finely reticulate, heterobrochate.
Habitus: shrub or tree to 9 m tall, sometimes scandent.
Habitat: forest, woodland, thicket, Acacia-Commiphora bush-
land, salt-flats and riverine vegetation; 0e700 (1200) m. At Kili-
manjaro in savanna woodlands, 900 m.
377
Distribution: K 7; T 2, 3, 6, 8; Malawi, Mozambique, Zimbabwe.
At Kilimanjaro rare.
Pollination: not kown; generally, species of Grewia L. appear to
be pollinated primarily by bees (Mawdsley and Sithole, 2010).
Triumfetta brachyceras K. Schum (Plate 21, 32e34). (Malvaceae
Grewioideae).
Grewia-type.
Shape: prolate e rhombic tall ~30  50 mm, circular.
Apertures: 3-colporate, rectangular broad e lalongate pori,
narrow colpi (4/5), constricted at equator.
Exine: tectate, finely reticulate, ± homobrochate.
Habitus: shrub (1-) 2e4 m high, sometimes spreading over a
wide area by horizontal stems.
Habitat: forest edges and clearings and swamps; 1200e2650 m.
At Kilimanjaro mainly in colline e submontane (riverine) forests
and grasslands, also in Chagga homegardens and lower montane
forest clearings; 900e2180 m.
Distribution: U 2e4; K 1, 3e6; T 1, 2, 4, 7; DR Congo, Rwanda,
Sudan, Ethiopia. At Kilimanjaro widespread.
Pollination: not known; for Triumfetta semitriloba flower visitors
are mainly solitary bee species, although social bees, beetles, flies,
bugs and butterflies have also been recorded (Collevatti et al.,
2000).
Triumfetta flavescens Hochst. ex A. Rich (Plate 21, 35e37).
(Malvaceae Grewioideae).
Grewia-type.
Shape: prolate e rectangular tall ~24  43 mm, circular.
Apertures: 3-colporate, rectangular broad pori, colpi (4/5) nar-
row, not constricted.
Exine: tectate, reticulate, ± homobrochate, lumina larger than in
T. brachyceras.
Habitus: small subshrub 0.35e2 m tall.
Habitat: Acacia-Commiphora and other dry bushland;
250e1600 m. At Kilimanjaro mainly in dry colline savanna wood-
lands and grasslands; 1000e1250 m.
Distribution: U 1; K 1e7; T 2, 3; Ethiopia, Somalia, Egypt, Saudi
Arabia. At Kilimanjaro widespread.
Pollination: not known; for Triumfetta semitriloba flower visitors
are mainly solitary bee species, although social bees, beetles, flies,
bugs and butterflies have also been recorded (Collevatti et al.,
2000).
5.2.76. Ulmaceae
Trees or shrubs with watery sap. A widely distributed family of
18 genera and ca. 150 species found in both tropical and temperate
regions, but best represent in the north temperate zone (Todzia,
1993).
5.2.76.1. Pollination. Flower morphology suggests wind pollina-
tion, however, several species have found to be anemophilous
(Todzia, 1993).
5.2.76.2. Pollen morphology. Based on exine sculpturing and
structure, pollen grains of the Ulmaceae can be placed into six
different types (Takahashi, 1989). Members of Ulmoideae all share
tetra- or pentaporate, oblate to spheroidal pollen grains varying in
size from 23 mm to 42 mm and distinguished by wholly granular
ectexine and rugulate sculpturing with spinules (Zavada, 1983).
Within the Celtidoideae five different types of pollen grains have
been described of which only Chaetacme, Trema and Celtis occur in
Tropical East Africa. Ampelocera (South America) pollen is distin-
guished by being periporate or tetra-to pentaporate and having a
palisade rather than granular ectexine (Takahashi, 1989). Chae-
tachme grains are similar to those of Ampelocera except that they
have a very thin endexine and a perforated exine with spinules. A
378 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
densely warty exine and a middle granular layer distinguished the
triporate pollen grains of Gironniera (SE Asia, Europe and South
America) (now Cannabaceae s.l.). Densely spaced warts and
microechinules and the absence of a granular layer in the exine
characterize Lozanella (South America), Parasponia (SE Asia), and
Trema (all now Cannabaceae s.l.). These grains are radially or
bilaterally symmetrical and either diporate or triporate. Apha-
nanthe (Central America, Madagascar, SE Asia), Celtis, and Pteroceltis
(Asia) (also all Cannabaceae s.l.) are tri-to pentaporate and all have
a middle granular layer in the extexine and spinules with a rod-like
substructure on the surface (Sattarian et al., 2006).
Celtis gomphophylla Engl (Plate 21, 38e40 & Plate 22, 1). SYN
C. durandii (Cannabaceae Celtidoideae).
Shape: circular, spheroidal, ~25 mm.
Apertures: 2, 3 or 4-porate with thick and protruding annulus,
pori circular to elliptic.
Exine: tectate, bacculate/granulate.
Habitus: much-branched deciduous tree, 5e25 m tall.
Habitat: lowland and upland rain-forest; 300e2000 m. At Kili-
manjaro mainly in colline and submontane riverine and Croton-
Calodendrum forests; 1080e1740 m.
Distribution: U 2e4; K 1, 4, 5, T 2, 3, 6, 7; Mozambique, Zambia
and Zimbabwe to the Cape Province of South Africa, also DR Congo
Republic and Angola to Nigeria and S. Tome . At Kilimanjaro
scattered.
Pollination: pollinated by insects, primarily by bees (Lemmens
et al., 2009).
Trema guineensis (Schum. & Thonn.) Ficalho (Plate 22, 2e3) SYN:
T. guineense (Cannabaceae Celtidoideae).
Shape: circular, spheroidal, ~24 mm.
Apertures: 3-porate, circular pori with protruding annulus.
Exine: psilate, scabrate.
Habitus: shrub, small or medium sized tree, up to 12 m tall.
Habitat: margins of lowland and upland rain-forest, often a
pioneer in clearings, also riverine forest; 0e2100 m. At Kilimanjaro
a pioneer tree in disturbed colline and submontane riverine and
Croton-Calodendrum forest, lower montane gorge forests, as well as
in Chagga homegardens; 780e1710 m.
Plate 22. ULMACEAE: 1 Celtis gomphophylla, 2e3 Trema guineensis, URTICACEAE: 4e5 Elatostemma paivaeanum, 6e7 Pilea johnstonii, 8e9 Urera hypselodendrum, VALERIANACEAE
(Caprifoliaceae): 10e12 Valeriana volkensii, VITACEAE: 13e15 Cissus oliveri, 16e19 Cyphostemma kilimandscharicum.
Distribution: U 1e4; K 1, 3e7; T 1, 8; Z; P; troughout Africa south
of the Sahara, Madagascar Is. and tropical Asia. At Kilimanjaro
widespread.
Pollination: not known for this species. Probably insect polli-
nated like Trema micrantha (Bawa et al., 1985).
5.2.77. Urticaceae
Herbs, shrubs, lianas or small trees. A family of 45 genera and ca.
1000 spp., widely distributed in the tropical and the temperate
regions (Friis, 1993).
5.2.77.1. Pollination. Only wind-pollination (Friis, 1993).
5.2.77.2. Pollen morphology. Grains of the Urticaceae are compar-
atively uniform, with 2e6 porate, (oblate-) suboblate-spheroidal,
small grains and an ± obscure exine sculpture and stratification.
Minute granules or echinae on the exine surface of about 0.5 mm or
less in diameter with a relatively low density. Pori circular in most
cases, with annulus (Erdtman, 1952; Sorsa and Huttunen, 1975).
Elatostemma paivaeanum Wedd (Plate 22, 4e5).
Shape: circular, spheroidal, ~12 mm.
Apertures: 2-porate, pori circular with annulus, annulus slightly
protruding.
Exine: intectate (?), very thin, scabrate.
Habitus: perennial herb. Creeping rhizomes end in one or a few
erect stems occasionally up to 1 m high, usually unbranched.
Habitat: lowland rain-forest or altitudinal transitional forest, in
the moist ground cover, often along streams; 900e2100 m. At
Kilimanjaro mainly in lower montane Newtonia gorge forests;
1170e2100 m.
Distribution: T 2, 3, 6, 7; widespread in the wetter parts of
tropical Africa, west to Guinee  south to Malawi. At Kilimanjaro
scattered.
Pollination: wind pollinated (Friis, 1993).
Pilea johnstonii Oliv (Plate 22, 6e7).
Shape: spheroidal to suboblate, circular, ~15 mm.
Apertures: 2-porate, pori circular with thin annulus which is not
protruding.
 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
Exine: intectate (?), thin exine, scattered micro-echinae.
Habitus: perennial herb with prostrate and ascending stems up
to 0.6 m high.
Habitat: in upland rain-forest and montane forest, sometimes as
an epiphyte; 1450e2900(-3600) m. At Kilimanjaro mainly in
middle montane Cassipourea and Ocotea forests and upper
montane Podocarpus, Juniperus and Hagenia forests; 1120e3200 m.
Distribution: U 2, 3; K 1, 3e7; T 2, 3, 7; Zaire (Ruwenzori), Sudan
(Imatong Mts.), S. Ethiopia, E. Zimbabwe. At Kilimanjaro
widespread.
Pollination: wind pollinated (Friis, 1993).
Urera hypselodendrum (A. Rich.) Wedd (Plate 22, 8e9).
Shape: spheroidal, circular; ~13 mm, however, dimensionally not
very stable.
Apertures: 3-porate, pori circular with thin annulus which is not
protruding.
Exine: intectate (?), thin exine, psilate.
Habitus: liana climbing to a height of 25 m or more.
Habitat: upland rain-forest and bamboo forest, especially in
clearings and near edges; (1050-) 1500e2350 m. At Kilimanjaro in
all colline to middle montane forest types and forest clearings;
850e2650 m.
Distribution: U 1e3; K 3e6; T 2, 3, 5e8; Sudan (Imatong Mts.),
Ethiopia, E. Zaire, Rwanda, Burundi, Malawi, E. Zimbabwe. At Kili-
manjaro widespread.
Pollination: wind pollinated (Friis, 1993).
5.2.78. Valerianaceae
Annual (occasionally biennial) or perennial herbs, rarely sub-
shrubs. A family of about 13 genera e only 4 genera, namely Cen-
tranthus, Fedia, Valeriana and Valerianella occurring in Africae and
about 400 species occurring mostly in the temperate zone of the
northern hemisphere. In East Africa, the few species known are
generally confined to high mountains (Kokwaro, 1968).
5.2.78.1. Pollination. Entomophilous (Willemstein, 1987).
5.2.78.2. Pollen morphology. Pollen grains 3 (4) colp(orid)ate,
suboblate-subprolate. Sexine usually thicker than nexine, tectate,
as a rule with small echinae (<3 mm). Colpi usually comparativley
short, their ends obtuse or acute. Pori very inconspicuous (Erdtman,
1952).
Valeriana volkensii Engl (Plate 22, 10e12). (Caprifoliaceae
Valerianoideae).
Shape: subprolate, trilobate, ~43  50 mm.
Apertures: 3 colp(orid)ate, colpi (5/6) wide at the equator with
thin margo and acute ends.
Exine: tectate, (micro) echinate.
Habit: perennial herb 5e10 (18) dm tall. Stems herbaceous,
usually clambering or erect.
Habitat: moist ground along stream sides, in moist bamboo
thickets, in upland rain-forest and in upland moor; 2300e3450 m.
At Kilimanjaro in upper montane Erica excelsa and Hagenia forests;
2860e3200 m.
Distribution: U 3; K 3, 5; T2; DR Congo Republic. At Kilimanjaro
rare, only on the northern slope.
Pollination: Valeriana is known to be pollinated by a variety of
insect species (e.g. Aluri and Robart, 1991; Waser and Ollerton,
2006).
5.2.79. Vitaceae
Woody climbers or vines, rarely small succulent trees. A pran-
tropical family of 14 genera and about 750 species, with a few
members in north temperate regions, and nine genera in E and SE
Asia (Wen, 2007).
379
5.2.79.1. Pollination. Insects and maybe wind (Willemstein, 1987;
Wen, 2007).
5.2.79.2. Pollen morphology. Pollen grains are 3-colporate, and vary
from oblate to prolate in shape. Amb often rounded triangular, then
angulaperturate. The sexine is reticulate and of about the same
thickness as the nexine (Erdtman, 1952). Dioecious species are
often found to have pollen dimorphy (e.g. Kevan et al., 1985).
Cissus oliveri (Engl.) Gilg (Plate 22, 13e15).
Shape: prolate, ~31  56 mm, trilobate?.
Apertures: 3-colporate, syncolpate, colpi narrow, pori circular
(Ø 7e8 mm) to elliptic broad with annulus.
Exine: tectate, thick exine, columellate (?), (micro) reticulate.
Habitus: climbing shrub 1.2e6 m long.
Habitat: forest, riverine forest, marshy areas, papyrus swamps,
wet grassland with scattered trees, sometimes on old termite
mounds; 930e2100 m. At Kilimanjaro in colline e lower montane
riverine and gorge forests, Croton-Calodendrum, Cassipourea and
Ocotea forests and clearings, as well as Chagga homegardens;
960e1930 m.
Distribution: U 2, 4; K 3e5, 7; T 2, 3, ?4, 6; Rwanda, Zaire,
Malawi, Mozambique and Angola. At Kilimanjaro widespread.
Pollination: by bees, flies, wasps, and butterflies (Martins, 2014).
Cyphostemma kilimandscharicum (Gilg) Desc. ex Wild & R.B.
Drumm. Agg (Plate 22, 16e19).
Shape: spheroidal, circular, ~35 mm.
Apertures: 3-colporate, large circular (Ø ~5e6 mm) to elliptic
broad pori with annulus, slightly wider than colpi, colpi (5/6)
tapering towards the poles with acute ends.
Exine: tectate, exine relatively thick, scabrate.
Habitus: robust ± herbaceous climber 3e20 m long.
Habitat: upland evergreen forest, mistforest, forest edges and
derived secondary scrub, bamboo thicket, streamside evergreen
bushland; 1400e2700 m. At Kilimanjaro mainly in submontane
Croton-Calodendrum forests and lower e middle montane Cassi-
pourea forests, as well as in Chagga homegardens; 1000e2780 m.
Distribution: U 1; K 1, 3e6; T 2, 3, 6, 7; Zaire, Sudan, Ethiopia,
Malawi and Zimbabwe. At Kilimanjaro widespread.
Pollination: by bees, flies, wasps, and butterflies (Martins, 2014).
6. Discussion and conclusion
With this pollen and spore atlas of tropical East Africa we add a
benchmark for the correct and consistent identification of pollen
grains and spores from East African sediment records and modern
pollen rain samples. We provide an overview on the range of most
important pollen and spore types found in the last glacial and
Holocene environmental archives in studies on and around Mt
Kilimanjaro. The additional information on plant pollination and
habitat makes this atlas a useful guide for palynological in-
vestigations, aiming at detailed and comprehensive reconstruction
of past vegetation, environmental and climate change in tropical
East Africa.
The selection of species is based on availability in the reference
collection as well as the ecological importance, abundance and
distribution of the species. In some cases, species are included
because the genus has significant abundance in tropical East Africa
(not only in the Kilimanjaro area) and the pollen morphology
within the genus does not show great variation (e.g Uebelinia).
Hence, the pollen morphology described here can help the identi-
fication of the genus, and potentially the species found in a
particular geographical area.
Knowledge on the pollen and spore flora of the region is still
limited, the list of pollen and spores is far from being exhaustive,
and additional investigations are required. We will also be adding
380 L. Schüler, A. Hemp / Quaternary International 425 (2016) 301e386
palynological descriptions of more pollen and spore types from
tropical East Africa as our reference collection is not exploited by far
and consistently growing. Further, all morphological descriptions of
pollen grains will be made accessible and searchable in the online
Go€ ttingen Pollen and Spore Image Database (http://gdvh.uni-
goettingen.de/).
Acknowledgments
We are very grateful to Anita Weißflog for compiling most of the
plates and helping with the literature research. We further thank
two anonymous reviewers for their constructive comments. Trav-
elling for the collection of pollen and spore material was mostly
funded by the DFG (German Research Foundation) within the
projects BE 2116/15-1 and SCHU 2978/1-2.
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