Journal of Vegetation Science 6: 777-786, 1995
© IAVS; Opulus Press Uppsala. Printed in Sweden
- Soil seed banks in dry Afromontane forests of Ethiopia -
Soil seed banks in dry Afromontane forests of Ethiopia
Demel Teketay1,2 & Granström, Anders1
1Swedish University of Agricultural Sciences, Faculty of Forestry, Department of Forest Vegetation Ecology,
S-901 83 Umeå, Sweden; Fax +46 90 166612; 2Permanent address: Alemaya University of Agriculture,
Faculty of Forestry, P. O. Box 138, Dire Dawa, Ethiopia; Tel. +251 5 111399
Abstract. The soil seed bank was investigated in four dry
Afromontane forests of Ethiopia. At least 167 plant species
were identified in the 0 - 9 cm soil layer with total densities
ranging between 12 300 and 24 000 seeds/m2. Herbs were
represented with the largest numbers of species and seeds in
the seed bank, while the contribution of tree species was
generally low. The overall vertical distribution of seeds was
similar at all sites with the highest densities occurring in the
upper three cm of soil and gradually decreasing densities with
increasing depth. Relatively high densities also occurred in the
litter layer. There were large differences in depth distribution
between species, suggesting differences in seed longevity. A
large number of species in dry Afromontane forests evidently
store quantities of seeds in the soil and this is in contrast to the
situation in most tropical rain forests, dry lowland forests and
savannas, where both the number of seeds and the number of
species are relatively small. It is possible that the strongly
seasonal and unpredictable climate of this region may have
selected for high levels of dormancy, and that herb regenera-
tion is associated with small scale disturbance. The fact that
most of the dominant tree species do not accumulate seeds in
the soil suggests that their regeneration from seed would be
unlikely if mature individuals disappeared. Most tree species
have relatively large seeds and poor long-distance dispersal;
this implies that restoration of Afromontane forests after de-
struction would be difficult. Since there is a diverse seed bank
of the ground flora, this component of the vegetation would
have a better chance of re-establishing. However, because
most cleared forest land is used for agricultural crop produc-
tion, it is probable that the seed bank will be depleted in only a
few years. Therefore, the future of the Afromontane forest
flora seems to depend on the successful conservation of the
few fragments of remaining natural forest.
Keywords: Floristic composition; Germination; Regenera-
tion; Soil seed density; Soil seed distribution.
Nomenclature: Cufodontis (1953-1972); Hedberg & Edwards
(1989); Friis (1992).
Introduction
The Afromontane Region extends intermittently at
altitudes above 2000 m from Sierra Leone in the west to
777
Somalia in the east, and from the Red Sea in the Sudan
Republic in the north to the Cape Peninsula in the south
(White 1983). A large portion of the Afromontane area
in Ethiopia was once covered by evergreen forests, but
only fragments remain and even these are at constant
risk of destruction. If the present trend of deforestation
continues, the remaining forests will disappear in the
very near future (Pohjonen & Pukkala 1990; Demel
1992; Tamrat 1993, 1994).
To manage the last remnants of Afromontane forests
and to make a future restoration programme possible, it
is essential to obtain information on the regeneration
ecology of at least the dominant species of the vegeta-
tion. Data on soil seed bank characteristics such as
species number, seed quantity and depth distribution
can provide insight into the regeneration ecology and
will in particular give an estimate of the regeneration
potential after disturbance (Thompson & Grime 1979;
Grime 1989; Whitmore 1983, 1991; Fenner 1985;
Garwood 1989; Thompson 1992). Studies in various
habitats have confirmed that, for species with a long-
lived seed bank, successful regeneration from buried
seeds usually occurs after disturbance (Bakker 1989;
Cavers & Benoit 1989; Keddy et al. 1989; van der Valk
& Pederson 1989). If there is no long-lived seed bank,
such regeneration takes place either through recently
dispersed seeds, or, in the case of undisturbed vegeta-
tion, through slow-growing seedlings and saplings.
Little is known about regeneration processes and
soil seed banks in dry Afromontane forests, and infor-
mation on fruiting phenology, dispersal, longevity, pre-
dation and germination ecology is almost lacking, even
for most of the dominant tree species. This lack of
knowledge of indigenous trees may have contributed to
the selection of Eucalyptus, Cupressus and other exotic
species for the few reforestation programmes which
have so far been undertaken (von Breitenbach 1961a, b;
Pohjonen & Pukkala 1990; Demel 1993; Tamrat 1993).
The present study is part of a research project deal-
ing with various aspects of regeneration and seed ecol-
ogy in dry Afromontane forests. Here, we present an
analysis of the soil seed bank at four widely separated
forest stands in the central and eastern highlands of
778 Demel Teketay & Granström, A.
Ethiopia, while concentrating on the question whether
there is a persistent soil seed bank in dry Afromontane
forests that could contribute to the restoration of natural
forest vegetation after severe disturbance.
Material and Methods
Study sites
Four study sites were selected in order to represent
the dry Afromontane forest vegetation found within the
altitudinal range of 2100 - 2700 m. They include ‘undif-
ferentiated Afromontane forests’ (Friis 1992). All stands
present show some evidence of human influence such as
selective tree cutting and livestock grazing, but they are
probably the best examples of natural vegetation avail-
able in the Afromontane area of Ethiopia today.
Munessa-Shashemene
The Munessa-Shashemene forest (hereafter referred
to as MS) is located about 200 km south of Addis Abeba
at 7° 13' N, 38° 37' E. The main forest blocks are found
on the escarpment and associated plateau lying between
the Rift Valley lakes and the eastern edge of the Rift
Valley. The bedrock is volcanic and soils are reddish,
freely draining and of medium to heavy texture (Lund-
gren 1971). The area is deeply incised by steep-sided
valleys cut by both permanent and seasonal water courses
and the altitude range extends from ca. 2100 m to 2700
m. The main wet season extends from July to October; a
less well defined rainy period occurs between March
and June. The mean annual rainfall in the forest is about
1250 mm. The dry season extends from November to
February. The mean annual temperature is 15 - 20 °C,
depending on elevation (Lundgren 1971; Chaffey 1980).
The natural forest is estimated to cover more than 15 000
ha. A detailed account on the recent history of the forest
is given by Chaffey (1980).
Gara Ades
The Gara Ades forest (GA) is situated ca. 400 km
east of Addis Abeba at ca. 9° 20' N, 41° 15' E. The
bedrock is volcanic and the soil is a dark grey loam with
a relatively high content of organic matter. Soils are
generally deep and contain many loose rocks up to about
15 cm in size. At the summit and on ridges the soil is
shallow and the rock is exposed in some places (Murphy
1968; Uhlig & Uhlig 1990). The altitude ranges from
2200 m to 2775 m. The mean annual rainfall is about
1200 mm with the highest rainfall extending between
June and September and a less pronounced wet period
occurring in March and April. The mean annual tem-
perature ranges from 15 - 20 °C (Anon. 1988). The forest
covers an area of ca. 500 ha. It was originally dominated
by Juniperus procera and Podocarpus falcatus, but
most of the larger Juniperus trees have been removed
through selective cutting since the 1950s. The floristic
composition of the forest has been reported by Uhlig &
Uhlig (1990) and human impact has been described
elsewhere (Demel 1992).
Menagesha
The Menagesha forest (ME) is situated 30 km south-
west of Addis Abeba at 9° 00' N, 38° 35' E. The area is
part of the fairly well preserved volcanic dome of Mt.
Wachacha (Mohr 1971). The soils are light brown to
reddish brown and shallow at the higher altitudes but
deeper at the lower altitudes (Tamrat 1993). The topog-
raphy of the area is extremely dissected, with alternating
ridges and valleys dominating the landscape. The forest
covers the northwestern and southwestern half of the
mountain with its lower boundary at ca. 2300 m and
extending up to ca. 3000 m, while the eastern slope has
been converted into farmland. There are no meteoro-
logical data available. The mean annual rainfall re-
corded at Holeta (2000 m), a town 5 - 7 km west of the
forest, is 1225 mm. There are six rainy months, April-
September, with most of the rain falling in July - Sep-
tember. The period between October and January is
relatively dry. The mean monthly temperatures range
from 12.4 °C to 15.9 °C (Tamrat 1993). The total area of
natural forest is 2720 ha (Forest Rangers, pers. comm.
1994). Detailed accounts on the history and floristic
composition of the forest can be found in von Breitenbach
& Koukol (1962), Gilbert (1970) and Sebsebe (1988)
while the phytosociological and structural attributes are
reported by Tamrat (1993).
Wof-Washa
The Wof-Washa forest (WW) is situated ca. 160 km
northeast of Addis Abeba at 9° 45' N, 39° 45' E. Bed-
rocks are volcanic (Chaffey 1979) and black soils are
predominant on the flat highlands as well as on the
bottom of wide valleys while reddish-brown and heavy
loams are predominant on the mountain and valley
slopes as well as on better drained areas (von Breitenbach
1961a, b). The forest occupies part of the eastern escarp-
ment of the northwest highlands. The escarpment is
broken up by a number of valleys and rivers which
penetrate the forest interior. The altitudinal range is
2100 - 3600 m. The very steep slopes and rugged topog-
raphy may have contributed much to its preservation
when adjacent forests were completely cleared. The
closest weather station is at Debre Berhan (2640 m), 30-
35 km west of the forest. Here, the average annual
rainfall is 800 mm and the average annual temperature is
15.8 °C (Daniel 1977). There are six rainy months from
 - Soil seed banks in dry Afromontane forests of Ethiopia -
March to April and from June to September, with very
high precipitation in July and August. The size of the
forest is difficult to ascertain; estimations range from
3500 - 10 000 ha (von Breitenbach 1962; Anon. 1991).
According to the inhabitants, there has not been any
commercial exploitation of the forest. The phyto-
sociology and structure of the forest have been de-
scribed by Tamrat (1993) while the floristic composi-
tion is put into relation with biodiversity issues by
Demel & Tamrat (1995).
Soil sampling and analysis
At each site 10 plots were selected for soil sampling in
the closed part of each forest in order to roughly cover the
altitudinal range of each site. The sample plots at each site
were always at least 100 m apart. At each plot an area of 15
cm × 15 cm was marked and four separate soil layers were
removed using a sharp knife and a spoon and put into cloth
bags; these layers include litter layer and three succes-
sively deeper mineral soil layers, each 3 cm thick. Sam-
pling at all sites was done in the second half of March, at
GA in 1991 and at the other three sites in 1992. Germina-
tion trials were started within 1 - 2 weeks after sampling.
The GA samples were incubated in a greenhouse in Umeå
(Sweden) set at a day/night fluctuating temperature re-
gime of 20/12 °C, which is a typical temperature range
between day and night at the soil surface within the GA
forest (Demel unpubl.). On sunny days in the summer,
temperature in the greenhouse reached up to 30 °C. The
samples received 12 h of light each day from Osram HQIT
400 W fluorescent tubes except during the summer when
there were long hours of daylight. The samples from the
other three sites were incubated in a lath-house at the
Alemaya University of Agriculture, Ethiopia (altitude
ca. 2000 m). Here, the samples were shielded from direct
sunlight, but exposed to the ambient temperature which
ranged between 15 - 34 °C during the day and 8 - 15 °C
during the night. Soil samples were in all cases spread to
a thickness of ca. 1 cm on cotton cloth in plastic trays and
kept continuously moist.
Seedlings started to emerge after about a week, and
those which were readily identifiable were recorded and
discarded. Those difficult to identify at the seedling
stage were transplanted and grown separately until they
could be identified. Sometimes this required flowering
of the plants. They were then pressed, and later identi-
fied at the Alemaya University of Agriculture Her-
barium or at the National Herbarium in Addis Abeba.
Every eight weeks, the soil samples were stirred to
stimulate seed germination.
After six months of incubation, the germination
trials were stopped and the samples were sieved using a
mesh size of 0.5 mm. All material retained on this mesh
779
was inspected for seeds, which were identified to spe-
cies if possible, using local reference material. Seed
viability was assessed by dissection; seeds were consid-
ered viable if their content was white and firm. To
determine the habitat requirements of the species recov-
ered from the soil seed bank, field observations were
made during trips in 1992 and 1993.
Data processing
Vertical distribution of seeds of a species is assumed
to reflect the longevity of its seeds in the soil (Leck
1989; Kjellsson 1992). To get a rough idea about the
ability of the species to survive in the soil, average depth
distribution was calculated for each species using the
following formula:
(SL × 0.5) + (ST × 2.5) + (SM × 5.5) + (SB × 8.5)
Average depth =
Total number of seeds
where SL = number of seeds in the litter layer;
ST = number of seeds in the first soil layer (0 - 3 cm);
SM = number of seeds in the second soil layer (3 - 6 cm);
SB = number of seeds in the third soil layer (6 - 9 cm).
The multipliers 0.5, 2.5, 5.5 and 8.5 are the depth (in cm)
from the litter surface to the middle of the litter layer,
first, second and third soil layers, respectively. The litter
layer varied in depth but in these calculations an average
depth of 0.5 cm has been assigned.
To compare similarities among sites, both in terms
of species composition and number of seeds, the data
were analysed using Principal Component Analysis with
the computer program SIMCA-S (Anon. 1992-1994). In
the analysis, only taxa identified to species level and
with at least 10 seeds recovered from the soil were
included. Species which were found to be outliers or
which did not contribute to the variation in the data were
excluded from the analysis. Sites were used as dummy
variables in the analysis. Before the analysis, the number
of seeds of each species in each sample was first con-
verted into a percentage value in relation to the highest
number of seeds of that species. The percentage values
were then transformed to the 1 - 9 scale similar to that of
van der Maarel (1979) as follows: < 1 % seed (s) = 1; 2
% = 2; 3 % = 3; 4 % = 4; 5 - 12.5 % = 5; 12.5 - 25 % = 6;
25 - 50 % = 7; 50 - 75 % = 8 and > 75 % = 9.
Results
Species composition and density of seeds in the soil
The total number of species recorded, excluding the
unidentified ones, was 92 from GA, 66 from ME, 62
780 Demel Teketay & Granström, A.

from WW and 58 from MS (App. 1). The total number Table 2. Soil seed density (seeds/m2) of species common to all
of species from the four sites was 167, of which 119 the sites and the five species with the highest soil seed density
were dicotyledonous herbs, 20 were shrubs, 12 grasses (*) at each site.
and/or sedges, nine climbers and seven trees (Table 1). Species Munessa- Gara Ades Menagesha Wof-
11 species occurred at all four sites (Table 2), 21 at three Shashemene Washa
sites, 35 at two sites and 100 at one site (App. 1). The
Crassula alsinoides 1556* 4222* 782* 2076*
most well represented family was Asteraceae with 18 Cyperus rotundus - - 960* -
species at GA, 14 species at ME, 12 species at WW and Dichrocephala integrifolia 329 1089* 142 338
11 species at MS. Five species of ferns indigenous to Droguetia iners 1440* 218 - 489
Erica arborea - - 1142* 5293*
Ethiopia were also recorded. Geranium simensis 22 84 138 80
The number of viable seeds in the soil samples (from Haplocarpha rueppellii - - - 2587*
both germination trial and sieving) correspond to a seed Helichrysum foetidum 76 1116* 9 116
Lobelia giberroa 2236* 218 422 -
bank density down to 9 cm in the soil of 12300/m2 at Monopsis stellarioides 440 147 1649* 484
ME, 13700/m2 at MS, 20100/m2 at GA and 24000/m2 at Oxalis corniculata 467 31 391 400
WW. Quantitatively, the seed bank was dominated by Pilea tetraphylla 1560* 53 49 -
Plantago palmata 369 - - 1382*
relatively few species (App. 1). The five species with Poa annua 169 4 27 1409*
the highest seed density contributed 57 % of the total P. leptoclada 31 200 262 404
estimated seed bank at MS, 51 % at GA, 42 % at ME and Rubus apetalus 36 391 18 84
Solanum luteum 1022* 106 - -
53 % at WW (Table 2). Erica arborea, Haplocarpha S. nigrum 22 9 18 27
rueppellii and Crassula alsinoides had the highest aver- Veronica abyssinica 236 1929* 578* 9
age number of seeds (App. 1); for instance, E. arborea V. javanica 49 1867* 418 -
had 5293 and 1142 seeds/m2 of seeds at WW and ME,
respectively. Herbs had the highest number of species
and seeds compared with other life forms (Table 1). Vertical distribution of seeds
Seeds of woody species made up only a minor compo-
nent of the seed bank at all sites (except for the shrub The vertical distribution of the seed bank was similar
Erica arborea) and very few trees had considerable at the four sites, with the highest densities in the upper
amounts of viable seeds in the soil, i.e. Croton macro- 3 cm of soil and gradually decreasing densities with
stachyus (average = 33 seeds/m2 for sites where its seeds increasing depth; relatively high densities also occurred
were recovered; App. 1), Juniperus procera (127 seeds/ in the thin litter layer (Fig. 1a). The number of species
m2), Nuxia congesta (271 seeds/m2) and Ficus sur (498 present showed a similar but less pronounced trend (Fig.
seeds/m2). 1b). There was more variation between species than
In the PCA ordination, the sites were clearly sepa- between sites in relation to the vertical distribution. For
rated along the first and third principal component axes. instance, seeds of some species were almost entirely
The first component separated GA and WW while the confined to the litter layer, e.g. Juniperus procera,
third component separated MS and ME. The first and Clematis hirsuta, Girardinia diversifolia and Pilea
second components, could not separate MS and ME, tetraphylla. On the other hand some species have seeds
indicating that the two sites are very similar to each which are well distributed in all soil layers, e.g. Laggera
other and more so than they are to GA and WW in their crispata, Lobelia giberroa, Crassula alsinoides, Ve-
species composition and quantity of seeds in the soil. ronica abyssinica and Poa leptoclada, while some oth-
ers had most seeds distributed in the deeper layers, e.g.
Indigofera rothii, Solanum nigrum and Eragrostis
schweinfurthii. The average depth of seeds of the differ-
Table 1. Number of species in the different life forms at the ent species gives an indication of their distribution in the
four sites. soil (Fig. 2, App. 1). There were some differences re-
Life form Number of species garding life forms, with herbs, grasses and sedges found
Munessa- Gara Ades Menagesha Wof-Washa
at greater depths than trees, shrubs and climbers (Fig. 2).
Shashemene As mentioned earlier, woody plants generally had
Trees 2 7 2 -
low seed numbers in the soil, but there were some
Shrubs 6 12 8 3 notable exceptions. Even though Ficus sur is repre-
Climbers 4 6 3 2 sented by only very few adult trees in the forest at GA, it
Herbs 44 59 47 53
Grasses and sedges 2 8 6 4
had the highest number of seeds (498 seeds/m2) among
the trees, and these seeds were well distributed in the
Total 58 92 66 62 deeper layers. Most seedlings of this species emerged
 - Soil seed banks in dry Afromontane forests of Ethiopia -
Fig. 1. Vertical distribution of seeds (a) and species (b) at the four sites (depth refers to the midpoints of the sampled layers).
from samples in the vicinity of mature, fruiting trees
(data not shown), but some seedlings also emerged from
samples taken in areas where mature Ficus trees were
absent. Of all species Erica arborea (ME and WW) had
the highest number of seeds, and the widest distribution,
both horizontally and vertically, especially in WW
(App. 1). Another woody species, Urera hypselodendron,
which is often found amongst climbing trees and shrubs
along road sides and in secondary vegetation, had large
numbers of seeds in the deeper layers at three of the
sites.
Soil sieving
Many seeds were recovered from all sites by sieving
the soil. These were categorized as dormant, eaten or
dead based on visual examination. The total number of
seeds recovered from the soil was 264 (representing five
species) from MS, 908 (more than 11 species) from GA,
639 (five species) from ME and 133 (five species) from
WW. Most seeds came from relatively large-seeded tree
species. The proportions of viable, eaten and dead seeds
of each species at each site are presented in Table 3.
Juniperus procera was the most common species with a
total of 65 viable seeds recovered, vs 21 seedlings that
had emerged in the germination trials. Closed but empty
seed coats, i.e. seeds that had rotted in the soil, were 14
× more numerous than viable seeds (data from germina-
tion trial and sieving combined). Seed coats with signs
of predation were about twice as numerous as viable
seeds. In the case of Podocarpus falcatus, no seedlings
had emerged in the trials; two out of 52 seeds sieved
from the soil were viable, 47 were empty, and three had
been eaten.
781
Discussion
Our results show that many plant species in dry
Afromontane forests store large quantities of seeds in
the soil, in contrast to the situation in most tropical rain
forests and savannas where both the number of seeds
and the number of species are relatively small (Keay
1960; Guevara & Gomez-Pompa 1972; Liew 1973; Hall
& Swaine 1980; Uhl et al. 1981; Enright 1985; Young
1985; Putz & Appanah 1987; Young et al. 1987; Saulei
& Swaine 1988; Garwood 1989; Alvarez-Buylla &
Martinez-Ramos 1990; Skoglund 1992). Another con-
trast between tropical rainforests and dry Afromontane
forests is that seed banks in many tropical rain forests
are dominated by pioneer species, particularly trees
(Thompson 1992), whereas most of the seed bank spe-
cies in the dry Afromontane forests are herbs which are
also well represented in the standing vegetation. Not
much is known on the life forms (sensu Raunkiaer 1934)
of the herbs and grasses found here, but it appears that a
large proportion of the species in the seed bank are
annuals (around 60 %, based on our own observations
and the literature; App. 1). Many of the herbs seem to
recruit after minor disturbances such as single tree wind
throw, grazing, and soil disturbance by wild pigs (own
observations), which are common phenomena in Afro-
montane forests. These disturbances in turn favour the
incorporation of seeds into the soil.
We have tentatively grouped species with more than
five seeds recovered (germination trial and sieving com-
bined) and an average depth distribution at or greater
than 1.5 cm as persistent seed bank species (App. 1);
these amount to 86 species, 81% of them being herbs.
We believe that these criteria are relatively conservative
and several species with small seed populations are
782 Demel Teketay & Granström, A.

Table 3. Apparently viable (VI), eaten (EA) and dead (DE) seeds sieved from soil samples after the germination experiments
terminated.

Species Munessa-Shashemene Gara Ades Menagesha Wof-Washa

VI EA DE VI EA DE VI EA DE VI EA DE

Trees
Croton macrostachyus 1 - 1 2 - 2 - 1 - - - -
Juniperus procera 3 - 90 45 141 534 17 42 549 - 26 40
Olea europaea subsp. cuspidata - - - 4 1 20 - 2 10 - - -
Podocarpus falcatus - 3 43 2 - 4 - - - - - -
Rhus sp. - - - - - 2 - - - - - -
Shrubs
Indigofera rothii - - - 14 - - - - - - - -
Rubus sp. - - 49 4 - 26 - - - - - 20
Climbers
Clematis sp. - - - - - 1 - - - - - -
Galium sp. - - - 3 - 6 - - - - - -
Herbs
Chenopodium sp. - - - 4 - 5 - - - - - -
Geranium sp. - - - 1 - - - - - - - -
Girardinia diversifolia 74 - - - - - - - - - - -
Trifolium sp. - - - - - - - - - 38 - -
Unidentified
Sp. A - - - - - - 1 1 2 - - -
Sp. B - - - - - - - - - 2 - -
Sp. C - - - - - - - - - 7 - -
Sp. D - - - - - - - 3 11 - - -
Others - - - 7 18 62 - - - - - -

excluded. A low-density seed population in the soil does
not by itself infer short persistence, but the limit of five
seeds was adopted as a reasonable sample size for the
depth distribution. We have also attempted to group
species into four categories with respect to their habit,
based on observations in the field and literature, i.e.
shade-intolerant/gap-colonizing species (G), indifferent
(establishing both in gaps and shade) (I), shade-tolerant
(found mainly under shade of trees and shrubs) (S), and
uncertain (no information about their preferences) (U).
The majority of the species with persistent seed banks
(58 %) are gap colonizers (App. 1). Several species, e.g.
Poa annua, Solanum nigrum, and Stellaria media, are
weeds and ruderal species of Eurasian origin which have
become cosmopolitan.
The data show that the majority of woody plant
species do not accumulate seeds in the soil. Previous
floristic studies of the standing vegetation at the four
sites show that there are at least 66 species of woody
plants, including woody climbers, at the Munessa-
Shashemene forest (Lundgren & Lundgren 1969), 58 at
Gara Ades forest (Uhlig & Uhlig 1990; pers. observ.),
55 at Menagesha forest (Sebsebe 1988) and 54 at Wof-
Washa forest (Demel & Tamrat 1995). In several in-
stances, soil samples taken virtually under the canopy of
large mature trees of many common species, e.g.
Podocarpus falcatus, Olea europaea ssp. cuspidata,
Hagenia abyssinica, Bersama abyssinica and Ekebergia
capensis, failed to reveal the presence of viable seeds.
This general lack of a soil seed bank in tree species may
be due to several factors. Some species, notably B.
abyssinica and E. capensis, appear to have recalcitrant
seeds, adapted to germination more or less immediately
after seed shed (Demel unpubl.). Others may have a
capacity to store seeds in the soil for several years. In
burial experiments 99 % and 93 % of the seed popu-
lations of Acacia abyssinica and Croton macrostachyus,
respectively, were still alive after 37 months of burial in
the soil (Demel unpubl.). It is likely that predation of
these large seeds on the soil surface prohibits the build-
up of a substantial seed bank. Pudden (1958) reported
from Kenya that large quantities of seed of Juniperus
procera were eaten by monkeys in the tree canopies,
and about 75 % of the seeds reaching the ground were
consumed by rodents.
Although we have no quantitative data on seed pre-
dation, inspection of rodent nests at GA revealed large
heaps of seed coats of both Olea europaea ssp. cuspidata
and Podocarpus falcatus. A considerable number of
eaten seeds of J. procera and a small number of seeds of
O. europaea ssp. cuspidata and P. falcatus (Table 3)
were also found in the soil which was sieved after the
germination trial was terminated, suggesting that preda-
tion may play an important role in reducing the seed
 - Soil seed banks in dry Afromontane forests of Ethiopia -
Fig. 2. Average vertical distribution of seeds of the 167
species grouped by life form.
populations of these species.
A considerable number of seedlings and saplings of
some tree species, e.g. B. abyssinica, E. capensis, P. fal-
catus and O. europaea ssp. cuspidata are found within
these forests (Demel unpubl.), suggesting a successful
seedling bank habit. This may be the typical strategy for
tree regeneration in the Afromontane forests.
Only some woody species show evidence of a seed
bank habit: Croton macrostachyus, Erica arborea, Ficus
sur, Hypericum revolutum, Rubus apetalus, Urera hyps-
elodendron, Nuxia congesta, Laggera crispata, Lobelia
giberroa, Solanecio gigas and Halleria lucida (App. 1).
Ficus species are also prominent in the soil seed banks
of other tropical forests (Hall & Swaine 1980; Hopkins
& Graham 1983; Graham & Hopkins 1990; Russell-
Smith & Leucas 1994). Most of the species (both herbs
and woody plants) which have been shown in the present
study to accumulate buried seeds in the soil are gap-
colonizing, and have small seeds ranging 0.5 - 2.0 mm in
length, which are sometimes capable of long-distance
dispersal. Seeds with the ability to stay viable for a long
time tend to be very small and selection for possession
783
of a persistent seed bank appears to favour smaller seeds
(Thompson 1987). This group of species has the follow-
ing attributes related to their success in establishing
after disturbance: production of large numbers of seeds,
means of long distance dispersal, formation of persist-
ent soil seed banks and the capacity to remain viable in
a dormant state for a long period of time (Whitmore
1991). Dormancy mechanisms for most of the gap-
colonizing species in the present study are unknown, but
for some, germination tests have been performed. For
example, there were requirements of both light and
fluctuating temperature for germination in fresh seeds
of three Veronica species and only fluctuating tempera-
ture for three Solanum species (Demel unpubl.). The
fact that many species have a seed bank deeply distrib-
uted in the soil (App. 1; Fig. 2) suggests that a large part
of the seed populations is quite old (Kjellson 1992), and
that dormancy is broken only in connection with distur-
bance.
The PCA revealed that 66 species, only 39 % of the
total species number, can explain the variation in the
soil seed bank data. This confirms that there is a strong
similarity among the four sites regarding their soil seed
bank composition. However, the variation caused by the
66 species was sufficient to split the sites along compo-
nents 1 and 3. Some species contributed much to the
separation of one or the other site. For instance, Droguetia
iners, Girardinia diversifolia, Pilea tetraphylla, Sola-
num luteum and Carduus nyassanus had more seeds in
MS than the other sites. Halleria lucida, Eragrostis tef,
Cynodon dactylon and Dichrocephala integrifolia have
been recovered only at GA. These differences have been
used in the PCA to explain the separation of sites.
However, it was not possible to make any ecological
interpretation based on the distribution of these species
on the plots.
Dry Afromontane forests can be characterized by
possessing large populations of buried seeds of herbs,
grasses and sedges whereas trees mainly have persistent
seedling banks and the ability to sprout from damaged
roots or shoots. The fact that most of the dominant tree
species do not accumulate seeds in the soil suggests that
the regeneration of primary tree species from seeds
would be prevented by removal of mature individuals.
Most species are relatively large seeded and apparently
have poor long-distance dispersal, suggesting that resto-
ration of Afromontane forests would be difficult and
slow to accomplish if they are destroyed. The ground
flora vegetation has a better chance of natural recovery,
because it has a very diverse seed bank, probably with
great seed longevity. However, since most cleared for-
est land is used for cereal production with annual culti-
vation, it is likely that the seed bank will be depleted in
only a few years. Therefore, the future of the Afro-
784 Demel Teketay & Granström, A.
montane forest flora seems to depend on the successful
conservation of the few remaining fragments of natural
forest.
Acknowledgements. Financial support for this project was
obtained from the Swedish International Development Agency
and the Alemaya University of Agriculture. Logistic support
was provided by the Faculty of Forestry, Faculty of Agricul-
ture and Debre-Zeit Research center (AUA), State Forest
Offices at the four sites, ACD and ETH. We thank Olle
Zackrisson, David Wardle and three anonymous reviewers for
comments on the manuscript, Anders Nordgren and Christer
Albano for their help in the multivariate analysis, Sylvia
Phillips for identifying grasses, Ove Ericsson for assisting in
data handling, Tamerat Abebe, Abdurazak Abdulahi, Mekdes
Mulu, Dawit Mulu and Keflie Lebassie for their help in the
field and lath-house.
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Received 7 February 1995;
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For App. 1, see p. 786.
786 Demel Teketay & Granström, A.

App. 1. Quantity of viable seeds in soil samples from the four sites (germination trials and soil sieving combined); species identified
at the species or genus level and having more than five seeds are included. S = seed bank type; P = persistent seed bank; H = habit;
G = gap-colonizing; I = indifferent; S = shade-tolerant; U = unknown. AD = average depth of occurrence in cm.

S H AD MS GA ME WW S H AD MS GA ME WW

Trees Geranium simensis P I 2.0 5 19 31 18

Croton macrostachyus P G 2.4 2 13 - - Girardinia diversifolia P G 2.0 203 - 3 -
Ficus sur P G 2.6 - 112 - - Gnaphalium declinatum P G 5.4 - - 93 -
Juniperus procera G 0.9 3 57 26 - Haplocarpha rueppellii P G 4.5 - - - 582
Nuxia congesta P G 3.9 - 61 - - Helichrysum foetidum P G 4.8 17 251 2 26
Olea europaea ssp. cuspidata S 1.3 - 8 1 - H. schimperi P G 2.5 - 2 3 31
Helichrysum sp. P G 4.6 - - 13 -
Shrubs Heracleum abyssinicum P I 1.7 - - - 92
Erica arborea P G 4.0 - - 257 1191 Hypericum peplidifolium P I 4.0 10 22 - 13
Halleria lucida P G 3.1 - 15 - - Hydrocotyle sp. P I 3.4 31 - - 187
Hypericum revolutum P G 2.1 - - - 89 Inula confertiflora P G 3.6 - 15 - 2
Indigofera rothii P G 6.2 - 16 - - Kalanchoe sp. P G 4.0 - - 13 -
Laggera crispata P G 4.2 193 50 12 - Lobelia neumannnii P U 5.4 - 128 - -
Lobelia giberroa P G 3.1 503 49 95 - Mimulis gracilis P I 5.3 - 29 - -
Maesa lanceolata P G 2.3 - 9 - - Monopsis stellarioides P I 4.9 99 33 371 109
Pentas schimperiana P G 4.3 - - 13 - Oxalis corniculata P I 4.2 105 7 88 90
Rhus glutinosa G 1.3 - 6 - - Phyllanthus fischeri P I 4.2 - 58 -
Rubus apetalus P G 3.3 8 88 4 19 Physalis peruviana P G 2.8 - 6 - -
Solanecio gigas P G 2.5 - - 35 - Pilea tetraphylla P I 1.5 351 12 11 -
Vernonia rueppeliana P G 4.2 - 7 - - Plantago palmata P I 2.7 83 - - 311
Vernonia sp. G 1.1 13 - - - Ranunculus oreophytus P G 4.9 - - - 5
R. stagnalis P G 3.0 - - - 116
Climbers Rumex bequaertii P G 5.0 1 4 - 18
Clematis hirsuta G 1.1 55 2 - 21 Sagina abyssinica P S 5.0 - - 1 22
Galium simense P I 4.1 - 159 3 - S. punctata P I 2.8 - 1 12 -
Urera hypselodendron P G 3.9 45 78 5 - S. simensis P I 4.0 - 7 2 83
Sida ternata P I 5.8 4 5 1 -
Herbs Solanum indicum P G 4.1 7 22 - -
Achryospermum schimperi P I 2.3 - 70 - - S. luteum P G 4.6 230 24 - -
Achyranthus aspera P G 2.1 5 5 4 - S. nigrum P G 4.9 5 2 4 6
Agrocharis incognita P G 4.4 - 8 - - Spilanthus mauritiana P I 4.2 10 62 - -
Alchemilla abyssinica P I 4.4 36 - 29 116 Stachys sidamoënsis P S 4.8 15 - - 6
A. cryptantha P I 3.3 1 16 - 39 Stellaria media P G 2.3 2 - - 14
Amaranthus sp. P G 5.5 1 - 8 - Tagetes minuta G 0.7 - 9 - -
Arenaria serpyllifolia P U 5.3 - 30 - - Trifolium sp. P G 4.7 - - - 6
Astrolinum adoensis P U 5.5 - 1 15 - Veronica abyssinica P I 4.3 53 434 130 2
Cardamine hirsuta P U 5.1 2 - 3 10 V. glandulosa P I 3.5 2 - - 13
Carduus leptacantha P G 4.0 40 - 5 - V. javanica P I 4.4 11 420 94 -
Cerastium afromontanum P I 2.1 9 - - 22 Viloa abyssinica P I 5.1 1 - - 216
C. octandrum P I 4.3 - 12 - -
Chenopodium murale P G 2.9 - 91 - - Grasses (Poaceae)
Cineraria abyssinica P G 4.0 - - 14 98 Cynodon dactylon P I 2.7 - 40 - -
Conyza abyssinica P G 6.5 - 50 - - Eragrostis schweinfurthii P G 5.5 - 31 - -
C. bonariensis P G 4.7 30 - 40 102 E. tef P G 6.4 - 42 - -
C. hypoleuca P G 3.8 - 18 - - Festuca sp. P G 4.7 - - - 154
C. steudelii P G 6.3 2 38 - - Poa annua P G 3.8 38 1 6 317
Conyza sp. P G 3.3 - - - 52 P. leptoclada P G 3.6 7 45 59 91
Crassula alsinoides P I 4.0 350 950 176 467
C. schimperi P S 4.0 - 1 3 30 Sedges (Cyperaceae)
Dichrocephala chrysanthemifolia P I 6.6 - 213 - - Cyperus rotundus P G 3.2 - - 216 -
D. integrifolia P I 3.9 74 245 32 76 C. sesquiflorus P G 4.4 - 23 36 8
Droguetia iners P I 4.1 324 49 - 110
Total number of species 58 92 66 62