Weatherall ThomasCR

Seed Dynamics and Seedling Survival in Mainland
Thicket of the Eastern Cape

By

Clayton Richard Weatherall-Thomas

Submitted in fulfilment of the requirements for the degree Magister Scientae in the
faculty of Science at the Nelson Mandela Metropolitan University

January 2009

Supervisor: Prof. E.E. Campbell

i
Acknowledgements
I would like to thank my supervisor Eileen Campbell whose support was always readily
given and appreciated. Without you Eileen, this project would not have been possible.

Acknowledgement is necessary for the National Research Foundation (NRF), as well the
Nelson Mandela Metropolitan University who provided financial support. The
Department of Economic Development and Environmental Affairs (DEDEA) of the
Eastern Cape Province, as well as the Eastern Cape Parks Board (ECPB) is thanked for
collection permits, and permission to sample on their property. Thanks must also go to
the management of Shamwari Game Reserve, the management of Pretoria Portland
Cement (PPC) Loerie limestone quarry, and the management of African Brick, the
owners of the farm Welbedachtsfontein.

A very big thank you is owed Richard Fenwick, Pierre Swart, Anusha Rajkaran, Merika
Louw and Claire Browning who assisted me in the field. Special acknowledgement of
Richard and Pierre is necessary, as they lost much sweat and blood digging holes
underneath the canopy of my beloved but thorny Thicket, as well as Merika who spent
many hours in the glasshouse with me. Megan du Toit is thanked for her assistance in
data entry. Nelia Garner is also thanked for her help in the herbarium, as well as her
encouragement. Gratitude is owed to Andrew Gouws for assistance with the shade
enclosures, as well as Andy Smith for putting up with my continuous enquiries.

Much appreciation is owed to all the members of the Botany Department, and especially
the secretary Marilyn Philander, for the many cups of coffee and laughs shared. This time
of my life would have been that much duller without you.

Claire Browning is thanked again for being an awesome housemate, and for all the fun
times, as well as the company during the long days of writing up. Thanks goes to all of
my friends, whose continuous questions reminded me that I was not finished. I wouldnt
trade you guys in for anything!
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The largest thank you goes to my parents, who supported me through the hard times with
much food and prayer. I owe you a big debt of love and gratitude.

To my Lord and Saviour Jesus Christ, author and finisher of my faith, who alone gave me
the strength and motivation to get through it all.
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Table of Contents
Acknowledgements .......................................................................................................... i
Table of Contents .......................................................................................................... iii
List of Tables ................................................................................................................. vi
List of Figures ..............................................................................................................viii
Abstract ........................................................................................................................... x
Chapter 1 Introduction to Research .......................................................................... 1
1.1 General Introduction .............................................................................................. 1
1.2 The Thicket Biome ................................................................................................ 4
1.2.1 Vegetation of Thicket ...................................................................................... 5
1.2.2 Environmental Characteristics ......................................................................... 7
1.2.3 Classification of Thicket ................................................................................. 8
1.2.4 Degradation and Transformation ................................................................... 14
1.2.5 Regeneration and Restoration ........................................................................ 17
1.3 Research Rationale .............................................................................................. 19
Chapter 2 The seed bank in Mainland Thicket types across a vegetation cover
gradient ........................................................................................................................ 20
2.1 Introduction ......................................................................................................... 20
2.1.1 Classification of seed banks .......................................................................... 21
2.1.2 Composition and distribution of a seed bank ................................................. 21
2.1.3 Factors affecting the seed bank in Thicket ..................................................... 23
2.2 Materials and Methods ......................................................................................... 26
2.2.1 Study sites..................................................................................................... 26
2.2.2 Field Sampling .............................................................................................. 30
2.2.3 Laboratory study ........................................................................................... 31
2.3 Results ................................................................................................................. 32
2.3.1 Aboveground Vegetation .............................................................................. 32
2.3.2 Composition of the seed bank ....................................................................... 41
2.4 Discussion ............................................................................................................... 47
2.4.1 Aboveground vegetation and classification ................................................... 48
iv
2.4.2 Seedlings and the microenvironment ............................................................. 50
2.4.3 Characteristics of seed bank .......................................................................... 52
2.4.4 Similarity between seed bank and aboveground vegetation ........................... 54
2.4.5 Implications for regeneration through seed bank and seedlings ...................... 55
2.4.6 Conclusion .................................................................................................... 57
Chapter 3 Dormancy and germination in Thicket species ...................................... 59
3.1 Introduction ......................................................................................................... 59
3.1.1 Seed Dormancy ............................................................................................. 60
3.1.2 Types of Seed Dormancy .............................................................................. 61
3.1.3 Factors affecting the germination of Thicket species ..................................... 64
3.3 Results ................................................................................................................. 70
3.4 Discussion ........................................................................................................... 74
3.4.1 Seed Coat Induced Dormancy ....................................................................... 75
3.4.2 Embryo Dormancy ........................................................................................ 77
3.4.3 Conclusion .................................................................................................... 79
Chapter 4 Response of seedlings to drought and shade .......................................... 81
4.1 Introduction ......................................................................................................... 81
4.1.1 Seedling response to drought and shade ........................................................ 81
4.1.2 Seedling growth and survival in Thicket........................................................ 83
4.2.1 Study species ................................................................................................ 86
4.2.2 Germination of seeds .................................................................................... 87
4.2.3 Experimental design ...................................................................................... 88
4.3 Results ................................................................................................................. 89
4.3.1 Effect of shade .................................................................................................. 89
4.3.2 Effect of water .................................................................................................. 90
4.3.3 Effect of species................................................................................................ 93
4.4 Discussion ........................................................................................................... 94
4.4.1 Overall growth responses .............................................................................. 94
4.4.2 Response to shade ......................................................................................... 95
v
4.4.3 Response to drought ...................................................................................... 97
4.4.4 Implications for seedling survival and regeneration in different microhabitats
.............................................................................................................................. 99
4.4.5 Conclusion .................................................................................................. 101
Chapter 5 Conclusion ............................................................................................. 103
5.1 Opportunities for future research........................................................................ 109
References .................................................................................................................. 111
Appendix 1 ................................................................................................................. 127
Appendix 2 ................................................................................................................. 133
Appendix 3 ................................................................................................................. 138
vi
List of Tables
Table 1 STEP classification of Thicket, with mean annual rainfall (MAR) and character
species of each structural type. ...................................................................................... 15
Table 2 Mean (n = 5) cover of species in aboveground vegetation with greater than 1%
cover. BT = Buffels Thicket; AT = Albany Thicket; ST = Sundays Thicket; GT =
Gamtoos Thicket. .......................................................................................................... 33
Table 3 F-values of environmental measures between the three microhabitats of the four
Thicket types. AT = Albany Thicket; BT = Buffels Thicket; GT = Gamtoos Thicket; ST
= Sundays Thicket. ........................................................................................................ 38
Table 4 The number and growth form (G) of seedlings collected along transects (n=5) at
each study site and no. of sites (no.) where seedlings of each species occurs. AT =
Albany Thicket; BT = Buffels Thicket; GT = Gamtoos Thicket; ST = Sundays Thicket.
(Wo = woody, He = herb, S = succulent, Gr = graminoid, Cl = climber, Ge = geophyte)40
Table 5 Species richness (d), species diversity (H) and species dominance (C) of the
aboveground vegetation of sampling sites. ..................................................................... 42
Table 6 Mean (n = 4) number of seeds per m
2
, Growth form (G) and dispersal mechanism
(DM) of the dominant species in the germinable seed bank of the three microsites of the
sampling sites. (CA = canopy, ED = edge, OP = open). (v = present in aboveground
vegetation, sb = present in seed bank). Small letters indicate significant differences
between vegetation types for seed abundance and species richness. ............................... 44
Table 7 F-values for differences in seed density and species richness of the germinable
seed bank between the four vegetation types and three microhabitats............................. 45
Table 8 Mean (n=4) species richness (d), diversity (H), dominance (C) of the germinable
seed bank of study sites, and Sorensons co-efficient of community (CC). AT = Albany
Thicket; BT = Buffels Thicket; GT = Gamtoos Thicket; ST = Sundays Thicket. (Can =
canopy; Ed = edge; Op = Open)..................................................................................... 47
Table 9 Ecological characteristics and distribution of study species. .............................. 67
Table 10 Seed source, mean size (n = 10), mean weight (n = 10*) and relevant treatments
of study species. (MP = Mpumalanga, SC = southern Cape, EC = Eastern Cape, ex hort =
horticultural stock). (treatments: MS = mechanical scarification, HW = hot water, CS =
cold stratification, Sa = sand, Sh = shade). ..................................................................... 69
vii
Table 11 Mean (n = 3) germination (% SD) of study species exposed to different
treatments. Small letters indicate treatments with significant differences from the control.
...................................................................................................................................... 73
Table 12 F-values for measured responses of seedlings of the four study species to
various shade and drought treamtments. ........................................................................ 89
viii
List of Figures
Figure 1 Processes required for the successful regeneration of Thicket. ........................... 3
Figure 2 Classification of Thicket. Taken from Everard (1987)...................................... 11
Figure 3 Geographical location of study sites according to Everards (1987) classification
of Thicket, and climate data of the nearest weather stations. .......................................... 27
Figure 4 Geographical location of study sites according to STEPs (Vlok et al. 2003)
structural vegetation units of Thicket. ............................................................................ 28
Figure 5 Geographical location of study sites according to STEPs (Vlok et al. 2003)
biogeographical units ofThicket. .................................................................................... 28
Figure 6 Detrended Correspodence Analysis (DCA) of study sites, using coding
according to Everard's (1987) classification. MKT = Mesic Kaffrarian Thicket; XKS =
Xeric Kaffrarian Thicket; MST = Mesic Succulent Thicket. (x-axis eigenvalue = 0.745,
y-axis eigenvalue = 0.404. Cumulative percentage variance = 19.8%, Sum of all
eigenvalues = 5.813)...................................................................................................... 36
Figure 7 Detrended Correspodence Analysis (DCA) of study sites, using coding
according to STEP's biogeographic classification (Vlok et al. 2003). AT = Albany
Thicket; BT = Buffels Thicket; GT = Gamtoos Thicket; ST = Sundays Thicket. Sites are
also coded for Thicket (T) and Valley Thicket (VT). DCA statistics as in Figure 6. ....... 36
Figure 8 Air temperature (a), soil temperatures (b), light (c) and litter depth (d) in three
environments (canopy, edge and opening) of study sites. AT = Albany Thicket; BT =
Buffels Thicket; GT = Gamtoos Thicket; ST = Sundays Thicket.................................... 39
Figure 9 The relative growth form composition of seedlings found at study sites. AT =
Albany Thicket; BT = Buffels Thicket; GT = Gamtoos Thicket; ST = Sundays Thicket. 42
Figure 10 Detrended Correspondence Analysis (DCA) of the germinable seed bank of the
three microsites of the study sites. Blues are Buffels Thicket, Reds are Albany Thicket,
Yellows are Sundays Thicket and Purples are Gamtoos Thicket. (Filled circles = canopy,
Hatched circles = edge, Empty circles = open)............................................................... 45
Figure 11 Relative growth form composition of the microenvironments of the germinable
seed bank of study sites. AT = Albany Thicket; BT = Buffels Thicket; GT = Gamtoos
Thicket; ST = Sundays Thicket. (CA = canopy; ED = edge; OP = open). ....................... 46
ix
Figure 12 Relative dispersal mechanism composition of the germinable seed bank of the
three microenvironments of the study sites. AT = Albany Thicket; BT = Buffels Thicket;
GT = Gamtoos Thicket; ST = Sundays Thicket. (CA = canopy; ED = edge; OP = open).
...................................................................................................................................... 46
Figure 13 Germination rates of study species. Con () = control, MS () = mechanical
scarification, HW () = hot water, CS () = cold stratification, Sa () = sand, Sh () =
shade. ............................................................................................................................ 72
Figure 14 Mean growth responses (a = stem diameter, b = stem height, c= leaf number, d
= leaf length) of Thicket species to various shade treatments in a glasshouse. Lower case
letters indicate significant differences within species. Bars indicate standard error of the
mean. ............................................................................................................................ 91
Figure 15 Mean growth responses (a= stem diameter, b = stem height, c = leaf number, d
= leaf length) of Thicket species to various water treatments in a glasshouse. Lower case
letters indicate significant differences within species. Bars indicate standard error of the
mean. ............................................................................................................................ 93

x
Abstract
Subtropical Thicket is the dominant vegetation biome in the Eastern Cape. Little
regeneration through seedlings has been observed in Thicket. This is of serious concern,
as Thicket has been degraded over large areas of its distribution. An understanding of the
seed dynamics is necessary for any restoration project. A number of possible causes for
the lack of seedlings were therefore investigated. Study sites were best classified
according to the biogeographical regions of the Subtropical Thicket Ecosystem Planning
(STEP) project. The seed bank of Thicket is dissimilar to the aboveground vegetation, as
it lacks woody species and is dominated by herbs and grasses, making little contribution
to the regeneration of Thicket species. An increase in canopy cover resulted in lower soil
and air temperatures, as well as light, and an increase in litter depth. Seedling abundance
and species richness increases with mean annual rainfall, with woody seedlings being
dominant in the higher rainfall areas, and succulent species in the lower rainfall regions.
Anemochorous species dominated the germinable seed bank, and zoochorous species
were more prevalent underneath the canopy, compared to open areas, an indication of
active seed dispersal. No germination was recorded in many Thicket species, and
dormancy mechanisms were lacking in many of the species that did germinate.
Immersion in hot water increased germination in the hard seeds of Acacia karroo,
whereas shade enhanced germination in small seeded Burchellia bubalina. The response
of seedlings of four Thicket species with varying distribution to shade and drought was
studied. Ptaeroxylon obliquum and Aloe africana are drought tolerant species, whereas
Burchellia bubalina and Rhus dentata were intolerant to low soil moisture. All four
species were able to grow in deep shade, although Rhus dentata would be classified as a
shade avoider, unlike the other three species that are shade tolerant. This results in the
four species having different regeneration niches. It was concluded that dispersal and the
lack of safe sites for seedlings were the main limitations to seedling establishment in
Mainland Thicket.

1
Chapter 1 Introduction to Research
1.1 General Introduction
In the last 150 years, humans have caused a change of land use on a global scale,
resulting in the loss and continual decline of both the species diversity and functioning of
ecosystems on this planet (Gurevitch et al. 2006). Services provided to human beings by
ecosystems, which are generally not obvious, include the stabilisation of landscapes,
protection of soils, the retention of soil moisture, and the storing and cycling of nutrients
and carbon (Gaston 2000). The management of ecosystems may lead to their conversion
from a natural to a transformed state. This is usually accompanied by a loss of function
and the provision of services (Lechmere-Oertel 2003). This landscape transformation and
degradation has occurred in the Thicket biome, the dominant vegetation of the Eastern
Cape (Stuart-Hill 1991, Stuart-Hill 1992).

Thicket is highly adapted to its landscape as it is able to maintain high levels of biomass
throughout the year (Aucamp & Tainton 1984), as well as store large amounts of carbon
(Mills & Fey 2004, Mills et al. 2005) compared to other vegetation types in similar
climatic conditions. Thicket is able to tolerate xeric conditions that it is found in due to its
ability to modify the landscape by forming bushclumps (Palmer 1990). These
bushclumps ameliorate the environment by providing a microclimate underneath the
canopy that is more moist, cooler, richer in nutrients, as well as having less compacted
soil (Watson 2002, Lechmere-Oertel 2003). There is also an abundance of succulent
species that use Crassulacean Acid Metabolism (CAM) to fix carbon, such as
Portulacaria afra Jacq., Crassula ovata (Mill.) Druce and Aloe ferox Mill. (Midgley &
Cowling 1993).

The Thicket biome has suffered some of the heaviest levels of degradation of the biomes
in South Africa (Vlok et al. 2003), especially in the more arid interior of its distribution
in the Eastern Cape. Although it historically supported an array of indigenous herbivores
(Kerley et al. 1995), the main source of destruction is incorrect land management
practises relating to domestic herbivory (Hoffman 1987, Stuart-Hill & Danckwerts 1988).
2
This degradation results in the transformation of the landscape, from a dense shrubland to
an open pseudo-savanna (Hoffman & Cowling 1990) that is less productive and has a
lower carrying capacity (Stuart-Hill & Danckwerts 1988).

Very little or no regeneration through seedlings has been observed in transformed and
degraded Thicket (Stuart-Hill & Danckwerts 1988), with seedlings even being rare in
intact Thicket (Midgley & Cowling 1993). Where seedlings occur, they are limited to the
microclimate underneath the bushclumps (La Cock 1992, Sigwela 2004). The low
number of seedlings will lead to a lack of regeneration through genets in disturbed
landscapes where the protective canopy has been removed (Aucamp 1978). It is therefore
important to discover the reasons for this paucity of seedlings in Thicket (Fig. 1), and
whether this lack of regeneration is consistent across the whole biome.

Is the lack of seedlings in Thicket a reflection of poor propagule production? This does
not seem the case, as seed production is plentiful (Sigwela 2004) and non-seasonal
(Castley 1992; Watson 2002). Many plant species in Thicket flower profusely (Midgley
& von Maltitz 1991). Transformation does not affect the flowering of Pappea capensis
Eckl. & Zeyh., a keystone species in arid forms of Thicket (Sigwela 2004). Although P.
capensis does not fruit regularly (as is typical for many Thicket species) (Watson 2002),
it is a mast seeder, with most individuals fruiting at the same time (Sigwela 2004).

Is seed dispersal occurring in Thicket? Many Thicket species are vertebrate-dispersed
(Castley 1992, Watson 2002). As most large mammals have been removed from the
landscape (Kerley et al. 1995), seeds dependant on these species may no longer be
dispersed into new environments. The replacement of indigenous herbivores with
domestic species has led to lower levels of dispersal, as goats destroy more seeds than
kudu and bushbuck (Sigwela 1999). However, as most seedlings are not found
underneath their parent (La Cock 1992, Sigwela 2004), dispersal by birds is most likely
still occurring. As birds generally prefer the cover of bushclumps (Bruton 1992), few
seeds will be dispersed into open areas (Hall 2000).

3

Figure 1 Processes required for the successful regeneration of Thicket.

Are granivores consuming all the seeds produced in Thicket before they get an
opportunity to germinate? The removal of many bird and mammalian predators may have
led to greater numbers of small mammals, especially granivores. This would be
detrimental to recruitment for many of the plant species, as greater levels of consumption
will lead to less seeds and seedlings (Stuart-Hill & Danckwerts 1988). However, Sigwela
(2004) found significantly lower numbers of rodents in transformed xeric forms of
Thicket, suggesting that granivory is not responsible for the lack of seedlings. Whether
this applies to the Thicket biome as a whole is unknown.

Do important species in the Thicket biome accumulate persistent seed banks? Most long-
lived tree species, similar to the majority of Thicket plants (Cowling 1984), do not form
seed banks (Garwood 1989). The seed bank of Dune Thicket (Pierce & Cowling 1991),
Spekboom Thicket and Spekboomveld (Sigwela 2004) lacks woody species and is
dominated by grasses and forbs.

Do present environmental conditions promote the germination of dominant Thicket
species? Most woody species in Thicket have a tropical and subtropical origin (Cowling
1984), where most seeds of tree species do not remain alive for an extended period in the
Dispersal?
Seed bank?
Germination? Seedling survival?
Flower, fruit and
seed production?
Regeneration
and
Revegetation
Granivory?
4
soil (Vzquez-Yanes & Orozco-Segovia 1993). Portulacaria afra seeds lose viability
after only a few months (Whiting 1991, Pierce & Cowling 1991). Sigwela (2004)
suggests that most Thicket species have recalcitrant seeds, as few are found in the seed
bank. As previously mentioned, most species are endozoochorous, and the removal of
indigenous herbivores may have led to the loss of the mechanism necessary for dormancy
breaking (Sigwela 1999). On the other hand, the adaptations necessary for survival in the
seed bank (hard seed coat, small round seeds) are similar for surviving passage though
the gut (Pakeman et al. 2002). It is therefore unclear whether most Thicket species have
seeds that are able to persist in the soil, or whether they germinate immediately.

Does germination take place in Thicket, but seedlings of woody canopy species fail to
survive and contribute to the regeneration of the vegetation? Recruitment through
seedlings is assumed to occur infrequently, only after very wet seasons (Stuart-Hill &
Danckwerts 1988). Lack of seedlings may be a result of them being targeted by selective
herbivores (Midgley & von Maltitz 1991). Even though Sigwela (2004) observed that
few seedlings were consumed by goats over a period of a year and this lead to substantial
survival, he found few saplings and hence poor recruitment into the canopy.

1.2 The Thicket Biome
The concept of a Thicket biome is relatively new, and was first formally proposed by
Low & Rebelo (1998), although regarding it as separate vegetation unit had been
suggested by earlier authors (Cowling 1984, Palmer 1990). Previously, it had been
classified as being of Karoo origin (Acocks 1975), or as Savanna due to the dominance of
phanaerophytes and hemicryptophytes (Rutherford & Westfall 1986), or as Forest as it is
not fire-prone and functionally similar (Midgley et al. 1997). It was even called the
missing biome of South Africa (Everard 1991). As a result, this vegetation has many
aliases, including Valley Bushveld (Acocks 1975, Aucamp & Tainton 1984), Subtropical
Transitional Thicket (Cowling 1984; Everard 1987) and Subtropical Thicket (Palmer
1990; Midgley et al. 1997; Vlok et al. 2003). Low & Rebelo (1998) suggested it is
assigned as a biome because, although it shares the dominance of phanaerophytes (woody
5
shrubs and trees) with Savanna, it lacks the co-dominance of hemicryptophytes (grasses)
(Vlok et al. 2003, Cowling et al. 2005). Environmentally it is differentiated from
Savanna due to the absence of fire (Vlok et al. 2003). It cannot be classed as Forest as it
lacks in height (Holmes & Cowling 1993) and in development of strata and it generally
replaces Forest at sites protected from fire where rainfall is too low to support Forest
(Lubke 1996).

1.2.1 Vegetation of Thicket
Thicket has a transitional nature and cannot easily be defined (Cowling et al. 2005).
However, it is its species composition and dominant growth forms, as well as ecological
processes that characterize it. According to the extended definition of Thicket, there are
1 558 species present in Thicket, of which 322 (20%) are endemic (Vlok et al. 2003).
Most of these endemics are succulent, but to a lesser extent geophytes, and belong to the
Asphodelaceae, Apocynaceae, Crassulaceae, Euphorbiaceae and Mesembryanthemaceae
families. Asteraceae is the largest family in Thicket, but interestingly none of the 63
recorded grass species are endemic (Vlok et al. 2003).

Structure
Structurally, Thicket can be classified as a dense shrubland to low Forest, which is often
impenetrable and lacks strata (Everard 1987, Lubke 1996). It has a low canopy (0.5
3 m) dominated by largely evergreen multi-stemmed shrubs and single-stemmed trees,
mostly members of the Celastraceae, Ebenaceae and Anacardiaceae families (Vlok et al.
2003). As well as woody species, the canopy may consist of large succulents such as
Portulacaria afra, Aloe spp., and Cotyledon spp. (Cowling 1984). A number of species
emerge from the canopy, such as the tree euphorbias, aloes and Cussonia species (Vlok &
Euston-Brown 2002). Herbs, geophytes and succulents occur underneath the canopy,
with grasses being rare (Vlok et al. 2003). Climbers and lianas are common (Cowling
1984). The clearings within the Thicket are dominated by dwarf shrubs of Cape or Karoo
origin, as well as grasses in more mesic areas (Vlok et al. 2003).

6

Floristics
The Thicket biome is floristically variable, ranging from short single-stemmed trees in a
matrix of spekboom (Portulacaria afra) as well as multi-stemmed shrubs in xeric inland
areas, to forms that are transitional to tall dry Forest with few succulents in areas
experiencing higher rainfall (Cowling 1984, Everard 1987, Lubke 1998). Thicket is
regarded as transitional as it occurs in the Eastern Cape, the meeting place of four
phytochoria (Cowling 1984), and neighbours six biomes (Lubke 1998), forming mosaic
vegetation types with them (Vlok et al. 2003). It is dominated by species of Tongaland-
Pondoland origin, with Karoo-Namib being co-dominant in drier forms and Afromontane
Region species in wetter forms (Cowling 1984).

The majority of canopy species are adapted to high levels of herbivory as they are
palatable (Stuart-Hill 1991). Many are spinescent, resprout after damage and regenerate
mostly through vegetative growth (Cowling 1984, Midgley 1991, Stuart-Hill 1992), with
the exception of arborescent succulents (Midgley & Cowling 1993, Midgley 1997).
However, most shrubs are sensitive to defoliation (Stuart-Hill 1991) with recovery
occurring relatively slowly. Long rest periods are required between heavy periods of
browsing (Stuart-Hill & Danckwerts 1988). Most species have fruits that are dispersed by
birds or mammals (Cowling 1984), mostly by endozoochory (Bruton 1992, Castley 1992,
Sigwela 2004). The seeds of many species remain viable after passing through the gut of
an animal (Sigwela 1999), and this may be required to break dormancy for germination to
be improved (Cowling et al. 1997). Fruit production is also mostly aseasonal, allowing
frugivores and granivores to persist in the community throughout the year (Castley 1992).

The trees are adapted to xeric conditions (Aucamp & Tainton 1984), with many species
having either small leaves or large sclerophyllous evergreen leaves (Cowling 1984) and
are slow growing (Holmes and Cowling 1993). Many have succulent stems and/or leaves,
allowing them to survive extended periods of drought (Aucamp & Tainton 1984). Few
deciduous species occur, such as Ptaeroxylon obliquum (Thunb.) Radlk. (Coates-
7
Palgrave 2002). However, many require shaded conditions for establishment and survival
(Holmes & Cowling 1993) and these will not regenerate in degraded Thicket.

Grasses, which may be C3 or C4 (Vlok et al. 2003), are common in more mesic regions,
but rare in xeric areas, due to them being shallow-rooted in a landscape that experiences
unreliable rainfall (Everard 1987, Stuart-Hill 1992). Even though grass cover may be low
in Thicket and their contribution to animal production limited, they are sweetveld species,
and are able to produce large amounts of herbage within a limited period (Aucamp 1976,
Aucamp & Tainton 1984). Valuable grasses in Thicket are Digitaria eriantha Steud.,
Setaria sphacelata (Schumach.) Moss, Sporobolus fimbriatus (Trin.) Nees., Eragrostis
curvula (Schrad.) Nees, Themeda triandra Forssk. and Panicum spp (Aucamp 1976).

Thicket has a rich succulent flora as a result of its ability to survive regular droughts.
They may be of karroid (Cowling 1984) or tropical (Hoffman & Cowling 1991, Cowling
et al. 2005) affinity. A wide range of succulent growth forms occur, including herbs,
shrubs, trees and lianas, as well as leaf and stem succulents (Cowling et al. 2005). They
may either dominate the landscape, or be rare, depending on rainfall (Everard 1987), and
may be some of the first species lost from the landscape through overexploitation
(Aucamp & Tainton 1984, Moolman & Cowling 1994).

As most species are either succulent or have sclerophyllous leaves, as well as little grass
production, Thicket is generally fireproof as little combustible material builds up
(Moolman & Cowling 1994). However, in the mosaic vegetation types (with Fynbos in
the east and Grassland and Savanna in the west) fire plays a role in preserving the
patchiness of Thicket (Cowling et al. 1997, Vlok et al. 2003), and is necessary to
maintain the full complement of Thicket species (Vlok & Euston-Brown 2002).

1.2.2 Environmental Characteristics
The Eastern Cape, where the majority of Thicket is distributed, experiences a warm
temperate climate with a midsummer deficit (Cowling 1984). There is a relatively large
8
rainfall range in Thicket, as it grows in environments where rainfall is as low as 200 mm,
up to a maximum of 1 050 mm. The absolute amount of rainfall in not only important,
but also the season in which it falls. The dominant rainfall pattern of the Eastern Cape is
bimodal, with peaks in spring and autumn, with aseasonal and erratic rainfall being a
strong correlate of Thicket distribution (Vlok et al. 2003). Throughout the biome, at least
20% of rain falls in winter (April-August) (Vlok et al. 2003). Rainfall is generally
unreliable, and prolonged droughts are regularly experienced (Cowling 1984).

Thicket seldom experiences extreme temperatures, with the lowest mean minima ranging
from 7.7C at the coast to 0.9C in the arid interior. Similarly, mean maxima are lowest
at the coast (26.3C) reaching 32.6C inland (Vlok et al. 2003). There is little Thicket
above 1 000 m above sea level (Marker 1991), and frost is rare (Lubke 1998).

In the past, Thicket was thought to be limited to nutrient-rich soil (Cowling 1984; Palmer
1991), but it was later found associated with all major geologies, including the infertile
quartzitic sandstone (Vlok et al. 2003). Thicket modifies the soils in which it occurs by
enriching it with litter that decomposes in the absence of fire (Lechmere-Oertel 2003).

1.2.3 Classification of Thicket
Thicket has been classified in a number of ways, due to its transitional nature. Acocks
(1975) called this vegetation Valley Bushveld, as it is mainly limited to the river valleys
of the east and south coasts of South Africa. These valleys are hot and receive less rain
than the intervening ridges, from 500 mm to 900 mm per annum (Acocks 1975). Valley
Bushveld extends from the Gouritz River in the west to the Tugela River in the north-east
and is sub-divided as follows:
1. Valley Bushveld proper, northern variation, which extends from the Tugela River
southwards as far as the Great Kei River. In this form trees and shrubs are
abundant. There are some tree succulents but, in general, there is a scarcity of
succulents and an abundance of Acanthaceae and grasses.
9
2. Valley Bushveld proper, southern variation, which extends from the Great Kei to
the Kabeljouws valley. This is a dense bushland community with less
undergrowth and more succulents. This type has what can be called succulent
forests with Euphorbia triangularis Desf., Euphorbia tetragona Haw. and Aloe
species dominating. Acacia karroo Hayne is a common component of this
variation.
3. Fish River scrub, in the Lower Great Fish River valley, is extremely dense when
undamaged. The semi-succulent thorny scrub has a canopy height of 2 m, with
Euphorbia bothae Lotsy and Goddijn. as the dominant succulent.
4. Addo bush, is similar to the Fish River scrub but has some other species. Acocks
(1953) regards this variation to be a derivative of Alexandria Forest.
5. Sundays River scrub, in the wide flat lower Sundays River valley. This type
merges with the Addo bush and is less succulent and taller than the Fish River
scrub. Euphorbia spp. are common, with E. ledienii A. Berger replacing
E. bothae and E. grandidens Haw. being the dominant tree euphorbia. Aloe spp.
are conspicuous: A. ferox Mill., A. africana Mill., A. speciosa Bak. and A.
pluridens Haw. all being common arborescent aloes.
6. Gouritz River scrub in the valleys of the Gouritz, Little Brak and Great Brak
Rivers. This type is similar to Sundays River scrub except that the euphorbias are
replaced entirely by small aloes.

As well as Valley Bushveld, two other veld types recognised by Acocks (1953) are
included as Thicket (Vlok et al. 2003). Spekboomveld is the term used for largely
succulent thickets dominated by the spekboom (Portulacaria afra). Noorsveld refers to
the low thicket dominated by soet noors (Euphorbia caerulescens Haw.) (Acocks 1975).
These veld types are found inland of Valley Bushveld. These three vegetation types
(Hoffman 1987), together with Dune Thicket, a vegetation type recognised by Lubke et
al. (1986) as less succulent and confined to low coastal regions, make up the Eastern
Cape Subtropical Thicket (ECST).

10
Cowling (1984) divided Subtropical Transitional Thicket into the orders: Kaffrarian
Thicket and Kaffrarian Succulent Thicket. Kaffrarian Thicket consists of non-succulent
Subtropical Transitional Thicket communities with strong affinities with the
Afromontane Forest flora, although Tongaland-Pondoland endemics and linking species
dominate the phytochorological spectrum. Structurally, Thicket is a closed shrubland to
low Forest dominated by evergreen, sclerophyllous trees and shrubs with a high cover of
stem spines and vines (Cowling 1984). Kaffrarian Succulent Thicket occurs in dry areas
and is recognised by a high proportion of succulents and a flora transitional from
Tongaland-Pondoland to Karoo-Namib (Cowling 1983). Growth-forms are diverse and
include leaf- and stem-succulent shrubs, trees and vines, arborescent rosette succulents,
large- and small-leaved sclerophyllous and orthophyllous shrubs, low trees and vines,
grasses, forbs, annuals and geophytes (Everard 1987).

Everard (1987) identified four types of Subtropical Transitional Thicket in the two orders
proposed by Cowling (1984) based on syntaxonomic and structural attributes (Fig. 2).
Kaffrarian Thicket was divided into Xeric Kaffrarian Thicket and Mesic Kaffrarian
Thicket. Xeric Kaffrarian Thicket was found associated with the estuaries of the Fish,
Kariega, Gamtoos and Kabeljouws Rivers (Everard 1987). It forms a low Forest with a
high proportion of trees forming a closed canopy of 4 to 6 m in height. Succulents form a
small proportion of the flora with most species being woody. Endemism is low but
species richness and diversity is high. Rainfall can occur any time of the year and ranges
from 450 to 600 mm yr
-1
. Indicator species are Dovyalis rhamnoides (Burch. ex D.C.)
Burch & Harv., Cassine peragua L., Aloe ciliaris Haw. var. ciliaris and Dracaena
aletriformis (Haw.) Bos (Everard 1987).

Mesic Kaffrarian Thicket forms a very dense thicket of mainly woody shrubs and trees
Everard 1987). It occurs in subtropical river valleys that receive more than 800 mm yr
-1

and was identified in the Buffalo River area. Succulents form a very small proportion
with a high number of forbs and graminoids. It forms the most species rich and diverse
Thicket in the Eastern Cape but has the lowest endemism. Indicator species are

11

Figure 2 Classification of Thicket. Taken from Everard (1987).

Burchellia bubalina (L.f.) Sims, Brachylaena elliptica (Thunb.) D.C., Apodytes dimidiata
E. Mey. ex Arn. subsp. dimidiata and Acacia karroo (Everard 1987).

Two suborders of Kaffrarian Succulent Thicket have been recognised: Xeric Succulent
Thicket and Mesic Succulent Thicket (Everard 1987). The Xeric Succulent Thicket of
the Fish and Sundays Rivers is a low, relatively sparse thicket with the shrub canopy
ranging from between 2 and 2.5 m in height and having an average total cover of 63%. It
has a low number of succulents and a high number of endemics. It predominates in the
hot, dry river basins where average annual rainfall ranges from 300 to 450 mm. Indicator
species are Grewia robusta Burch., Asparagus striatus L.f., Brachylaena ilicifolia (Lam.)
E. Philips. & Schweick. and Euphorbia caerulescens (Everard 1987).

12
Mesic Succulent Thicket is an exceptionally dense, impenetrable thicket occurring in the
coastal areas from roughly the Sundays River mouth to the Gamtoos River valley
(Everard 1987). It has a total cover of 90% and an average height of 2.5 m and is
characterised by a high proportion of spinescent shrubs and woody creepers but also has
many succulents. Diversity and endemism is high and rainfall varies between 450 to
550 mm yr
-1
, falling mainly in the winter months. Indicator species include Aloe
africana, Sideroxylon inerme L. subsp. inerme, Aloe ferox and Euphorbia ledienii
(Everard 1987).

Low & Rebelo (1998) classify Valley Bushveld into five vegetation types within the
Thicket Biome on the basis of their distribution and the degree of succulence in the shrub
and tree species: Dune Thicket, Valley Thicket, Xeric Succulent Thicket, Mesic
Succulent Thicket and Spekboom Succulent Thicket.

Dune Thicket was identified by Everard as part of Xeric Kaffrarian Thicket but was
separated into its own vegetation type by Low & Rebelo (1998). It is a dense Thicket that
occurs at protected sites behind the foredune and at low altitudes (less than 30 m), and is
distributed along the coast from the Western Cape to KwaZulu-Natal (Lubke 1998).
Rainfall ranges from 900 to 1500 mm
-1
. The flora has affinities with that of the
Tongaland-Pondoland region. Dominant trees and shrubs include Mimusops caffra E.
Mey. ex A.D.C., Brachylaena discolor D.C., Strelitzia nicolai Regel & Krn, Sideroxylon
inerme, Maytenus heterophylla, Rhus crenata Thunb., Maytenus procumbens (L.f.) Loes.,
Passerina rigida Wikstr., Carissa bispinosa (L.) Desf. ex Brenan, Scutia myrtina
(Burm.f.) Kurz and Chrysanthemoides monilifera (L.) Norl. Liana and climbers are
common but there is little herbaceous cover.

Valley Thicket, also known as Valley Bushveld and Xeric Kaffrarian Thicket, consists of
a dense stand of woody shrubs and trees that dominate the river valleys from the eastern
parts of the Western Cape to KwaZulu-Natal (Lubke 1998). Rainfall ranges between 400
and 800 mm per year. The affinities of the flora is transitional between Tongaland-
Pondoland and Afromontane species. The closed canopy consists predominantly of
13
woody evergreen shrubs reaches up to 6 m in height, and succulent trees or shrubs being
less common. Many local endemic species occur, such as in the Albany hot spot.
Common species are Mystroxylon aethiopicum (Thunb.) Loes. subsp. aethiopicum,
Asparagus spp., Plumbago auriculata Lam., Dovyalis spp., Diospyros dichrophylla
(Gand.) De Winter, Euphorbia triangularis and E. tetragona (Lubke 1998).

Xeric Succulent Thicket occurs mainly in the Fish and Sundays River Valleys with
rainfall in these hot, dry valley basins between 300 and 450 mm per year (Lubke 1998).
Succulents are very common and the flora is transitional from Tongaland-Pondoland to
Karoo-Namibian. Its structure and species composition is that same as that of Everards
(1987) Xeric Succulent Thicket. Characteristic woody species are Grewia robusta,
Brachylaena ilicifolia, Gymnosporia capitata (E.Mey. ex Sond.) Loes. and Lycium
amoenum Dammer. Succulent species include Euphorbia caerulescens, Portulacaria
afra and Euphorbia bothae. There are few herbs, small shrubs and grasses such as
Pentzia incana (Thunb.) Kuntze, Chrysocoma ciliata L. and Cynodon dactylon (Lubke
1998).

Mesic Succulent Thicket is an exceptionally dense, impenetrable thicket occurring along
the coastal strip from the Sundays River mouth to the Gamtoos River valley (Lubke
1998). Rainfall varies from 450 to 550 mm per year and occurs mainly in the winter or
spring-autumn months. The description of this vegetation type follows that of Everards
(1987) Mesic Succulent Thicket.

Spekboom Succulent Thicket is found on the steep mountain slopes in the Eastern Cape,
as well as the eastern parts of the Western Cape (Lubke 1998). It is dominated by
Portulacaria afra that is also common in Valley and Succulent Thicket types - it merges
into these in river valleys. It occurs in a belt of about 400 to 1 060 m above sea level and
rainfall is 250 to 300 mm per year, falling mainly in autumn and spring. Other woody
species include Crassula ovata (Mill.) Druce, Lycium oxycarpum Dunal, Pappea
capensis, Euclea undulata Thunb., Rhigozum obovatum Burch., Grewia robusta, Rhus
14
spp. and Schotia afra (L.) Thunb. Many smaller Crassula spp. as well as succulent herbs
and grasses also occur (Lubke 1998).

In an attempt to classify thicket land classes for conservation planning, Vlok et al. (2003)
identified 112 Subtropical Thicket vegetation types falling within the Thicket biome.
This hierarchical classification system consists of four tiers, based on geography,
floristics, structure and grain (solid vs. mosaic). The first division is the geographical
separation of Dune and Mainland Thicket, with both of these sections divided into
biogeographical regions. Dune Thicket was defined along a west-east gradient, whereas
the Mainland Thicket was divided based on the main river drainage basins (Gouritz
Dune, Algoa Dune, Albany Dune, Transfish Dune, Gouritz, Gamka, Groot, Gamtoos,
Sundays, Albany, Fish, Buffels, Kei and Escarpment). The third tier consists of dividing
each drainage system into three structural types (Thicket, Valley Thicket and Arid
Thicket) based on the dominance of growth forms, as well as spinescence and lianas
(Table 1). Each structural type is further divided into two grain types based on whether
Thicket is occurs as a solid formation, or as a mosaic with another biome (fourth tier). Of
the 112 vegetation types, 78 are mosaics. The solid Thicket is further divided into
whether spekboom (Portulacaria afra) is dominant or subdominant. This classification
resulted in an expanded Subtropical Thicket concept and increased its extent in the study
area by between 1.8 and 2.8 times that of earlier assessments (Everard 1987; Low and
Rebelo 1998 respectively).

1.2.4 Degradation and Transformation
The dominant forms of vegetation disturbance in Thicket are shrub death (Stuart-Hill &
Danckwerts 1988) and herbivory, both domestic (La Cock et al. 1990, Moolman &
Cowling 1994) and indigenous (Stuart-Hill 1992, Kerley et al. 1995). The natural
disturbance regime would have been one of low disturbance, as most Thicket species
regenerate slowly (Hoffman & Everard 1987). The large herbivores that dominated the
landscape in pre-colonial times would have been responsible for gap-formation in
Thicket through browsing and path formation (Stuart-Hill 1992). With the removal of

15
Table 1 STEP classification of Thicket, with mean annual rainfall (MAR) and character species of
each structural type.
Structural Type MAR (SD) Character Species
Dune 649 (143) Sideroxylon inerme, Ischyrolepis eleocharis

Thicket 586 (129) Euphorbia triangularis, Olea europaea subsp.
africana, Scutia myrtina, Rhus pallens.

Valley 456 (100) Pappea capensis, Azima tetracantha, Rhus longispina,
Digitaria eriantha, Euphorbia spp., Aloe africana,
Crassula spp., Haworthia spp.

Arid 296 (94) Portulacaria afra, Euclea undulata, Euphorbia spp.
(non-arborescent forms)

megaherbivores, Thicket would become more dense, removing the regeneration niche of
shade-intolerant species of savanna origin (Holmes & Cowling 1993), as well as the
habitat of many rare and endemic succulents and geophytes (Cowling & Kerley 1995).
This densification is known as post-megaherbivore extirpation relaxation (Kamineth
2004). However, stocking goats at similar densities to indigenous herbivores results in
excessive shrub death and the opening of the canopy (Stuart-Hill 1992).

The difference between domestic and indigenous herbivores lies in the manner they
defoliate a plant, especially Portulacaria afra (Stuart-Hill 1991). Elephants, the historical
dominant defoliating agent in Thicket (Stuart-Hill & Danckwerts 1988) browse the top of
bushclumps, whereas goats, presently the largest cause of defoliation over large areas of
the biome, browse from the side. The top-heavy browsing does not prevent the plant from
forming a skirt of side-branches that have rooted, thus preserving the structural integrity
of the bushclump, and the microsite underneath it (Cowling & Kerley 1995). It promotes
coppicing and/or suckering from lateral roots (Stuart-Hill 1991). On the other hand, goats
and other smaller mammals defoliate the side branches, preventing the establishment of
new individuals, and destroying the ameliorated microclimate underneath the canopy
(Stuart-Hill 1992). However, the overstocking of elephants also greatly damages the
vegetation, leading to large decreases in cover and biomass (Cowling & Kerley 1995), as
well as the loss of species richness (Moolman & Cowling 1994). Therefore it is not
16
necessarily the type of animal that is important, but rather their stocking density (Stuart-
Hill & Danckwerts 1988).

Even though Thicket is a stable vegetation type and is able to resist most disturbances
(Cowling 1984, Everard 1987), large areas have been degraded or transformed through
unsustainable land practises and require rehabilitation (Hoffman & Cowling 1990). This
is compounded by different land-use systems, such as free-hold, state-owned and
communal tenure, which all cause different types and levels of degradation (Evans et al.
1997; Birch et al. 1999, Fabricius et al. 2002). The main sources of degradation are
overgrazing by domestic herbivory (especially goats) as well as the expansion of both
rural and urban settlements. In the higher rainfall areas, clearing of Thicket for cultivation
and dairy farming is an important impact (Palmer et al. 2004). Lesser threats include
mining, invasive species (both alien and karroid shrubs), fuel-wood and medicinal plant
collecting, fire, fragmentation and clearing along fence lines (Palmer et al. 2004).

Degradation of the vegetation results in loss of species, especially palatable ones
(Aucamp & Tainton 1984, Stuart-Hill & Danckwerts 1988, Stuart-Hill 1991 Fabricius et
al. 2002, Sigwela 2004) as well as the threatened species of succulents and geophytes
(Moolman & Cowling 1994, Lombard et al. 2001), and the invasion of alien species
(Aucamp & Tainton 1984). Structurally, there is a loss of biomass and cover (Penzhorn et
al. 1978, Stuart-Hill 1991, Cowling & Kerley 1995, Lechmere-Oertel 2003), and a
change in dominant growth forms from succulents and woody species to ephemeral
grasses and herbs (Sigwela 2004). Degradation and transformation is accompanied by
greater levels of soil erosion (Hoffman 1987, Kerley et al. 1999), the loss of soil nutrients
and carbon storage (Kerley et al. 1999, Mills & Fey 2004, Mills et al. 2005), as well as
lower levels of mycorrhizal fungi that promote nutrient absorption (Allsopp et al. 1996).

Thicket responds to degradation in two different patterns, depending on rainfall (Palmer
et al. 2004). Degradation in mesic forms of Thicket is accompanied by an increase in the
cover of woody species such as Acacia karroo, Lycium oxycarpum, Rhus undulata Jacq.,
Gymnosporia polyacantha (Sond.) Szyszyl. and Scutia myrtina mainly as a result of
17
excess domestic herbivory (Palmer et al. 2004). The encroachment of less desirable
woody species at the expense of open areas dominated by grasses may be due to a
changed fire regime, the removal of grass biomass by domestic herbivory, or the
competitive advantage of C3 shrubs over C4 grasses at increased CO
2
levels (Palmer et
al. 2004). Thus the landscape is transformed from a mosaic of bush clumps in a matrix of
grasslands, which supports hot but irregular fires, to a mixture of individual specimens of
Thicket species in a messy matrix of woody species, many of them invasive aliens,
with a grassy understorey that experiences cool but regular fires (Vlok & Euston-Brown
2002). Similar vegetation occurs when Thicket is completely cleared, either for
agriculture (Plamer et al. 2004) or rehabilitation after mining (Hall 2003).

In Xeric Thicket, overgrazing results in the loss of desirable woody and succulent species
(Palmer et al. 2004). In an untransformed state, the vegetation is dominated by long-lived
woody and succulent species in uninterrupted bushclumps that act as nutrient sinks and
are therefore high quality resource patches (Fabricius 1997). These bushclumps provide
cover and protection from the severe temperatures and light conditions and act as a
microhabitat for many plant and animal species that would not occur in the area without
this protection. When Thicket is overgrazed, it becomes less dense and there are larger
open areas that experience greater environmental fluctuations, as well as a loss of soil and
litter nutrients. After transformation the vegetation seems to support a different suite of
species. Mesic-adapted plant species are replaced by xeric-adapted species, as well as a
general change from palatable to unpalatable species (Fabricius 1997). The landscape is
transformed into a dwarf scrub desert, consisting of single umbrella-shaped trees in a
matrix dominated by annual herbs and grasses, or a pseudo-savanna (Stuart-Hill 1991).
These canopy trees will finally die off, leaving an alternate stable state consisting of a
highly eroded landscape with only ephemeral cover (Lechmere-Oertel 2003).

1.2.5 Regeneration and Restoration
Most species in Thicket have the ability to resprout after damage (Aucamp & Tainton
1984, Midgley 1991) with vegetative reproduction being the dominant form of
18
regeneration observed in Thicket. Seedlings are scarce, occur at low densities (Midgley &
Cowling 1993), and are limited to the microclimate underneath the canopy (La Cock
1992, Sigwela 2004). Seedling survival is low, even in rehabilitation patches consisting
of brushpiles (Todkill 2001). Sigwela (2004) states that seedling survival is high, with
few lost to herbivory. However, overall seedling survival decreased over time, as more
seedlings were missing due to mortality than new ones discovered (Sigwela 2004).

The regeneration of Thicket at a landscape level will begin with the formation of
bushclumps by the dispersal of seeds of pioneer species such as Azima tetracantha Lam.,
Gymnosporia spp., Lycium spp., Rhus spp. by birds, or by wind (Brachylaena spp.,
Tarchonanthus camphoratus L., Ptaeroxylon obliquum) (Vlok & Euston-Brown 2002).
There is evidence that there is a specific guild of pioneer species in Thicket that are
capable of forming a dense spiny tangle of recurved branches, casting a deep shade. This
promotes the establishment of shade-tolerant species, as well as preventing grasses and
fire from penetrating into the bushclumps. Bushclumps then expand and consolidate,
facilitated by spinescent lianas such as Asparagus spp. (Vlok & Euston-Brown 2002). In
higher rainfall areas, Thicket establishment may be initiated by non-spinescent
resprouting species such as Tecoma capensis (Thunb.) Lindl.and Plumbago auriculata,
although Acacia karroo may also be important (Hall 2003).

However, little spontaneous natural regeneration of Thicket has been observed, especially
of palatable shrubs, making restoration a difficult task (Aucamp & Tainton 1984). The
first step would be to restore the quality of the soil (Mills & Fey 2004), followed by the
creation of canopy cover. Restoration of Thicket has recently been attempted by planting
of P. afra truncheons (Powell et al. 2006). This is aimed at restoring carbon into the soil,
as well as introducing nutrients through litter (Sigwela et al. 2003, Mills et al. 2005).
With some cover established, birds should naturally disperse indigenous species back into
the landscape, where the ameliorated microclimate underneath the canopy will promote
seedling establishment and survival (La Cock 1992, Bruton 1999, Watson 2002, Sigwela
2004). However, fewer species are generally found underneath the canopy of P. afra
compared to other Thicket species (La Cock 1992) and active restoration of canopy
19
woody species, as well as the understory geophytes and succulents may be necessary for
the complete restoration of the landscape (Powell et al. 2006). In mesic Thickets,
restoration may occur naturally over an extended period of time, with the establishment
of species such as A. karroo (Vlok & Euston-Brown 2002), but once again active
introductions of the endemic succulents and geophytes may be necessary, as they have
limited dispersal distances (Hall 2003). Furthermore, to fully restore Thicket to its natural
state, indigenous herbivores, including the megaherbivores, need to be returned to the
landscape at sustainable levels, to restore the natural disturbance regime in Thicket, and
recreate the previous patchy state (Kerley et al.1999).
1.3 Research Rationale
Most of the research concerning seed dynamics and the regeneration of Thicket has
focused on xeric areas (Sigwela 2004), as well as Dune Thicket (Pierce & Cowling
1991). The aim of this research is to further the understanding of the seedling dynamics
of Thicket, focusing on the more mesic vegetation types (MAR > 400 mm).

In Chapter 2, the germinable seed bank in Thicket is investigated. The composition of the
seed bank is determined and whether it can act as a source of propagules for the
regeneration and restoration of Thicket. The question of why seedlings are limited to
Thicket clumps is explored, and if it could be the lack of dispersal into openings in
Thicket that prevents seedling establishment in interclump areas. A seedling survey was
included to illucidate the distribution of seedlings in Thicket.

Chapter 3 investigates the germination requirements of characteristic Thicket species.
The germination and dormancy mechanisms of species with hard seeds and thick seed
coats are contrasted with those that have thin seed coats, and this information is used to
predict their regeneration niche.

The drought and shade tolerance of four Thicket species are examined in Chapter 4. The
growth responses of three woody species are contrasted with the succulent Aloe africana.
The tolerance ranges of the four species are related to their possible regeneration niche.
20
Chapter 2 The seed bank in Mainland Thicket types across a
vegetation cover gradient

2.1 Introduction
Seed banks are important to the long-term maintenance and growth of plant species
(Louda 1989, Auld & Dunham 2001). They allow species to remain at a site after a large-
scale disturbance that results in the complete eradication of the aboveground population
(Adams et al. 2005), removing any source of seed for regeneration (Cummins & Miller
2002, Baskin & Baskin 1998). Seed banks allow species to spread the chance of
successful germination over time and space (Fenner & Thompson 2005). In semi-arid and
arid communities that experience long-term disturbances such as drought, species that
accumulate seed banks are able persist in the landscape (Kinloch & Friedel 2005b) and
respond quickly to rainfall events (Kinloch & Friedel 2005a). Seed banks are
characteristic of species in communities that experience large, irregular disturbances
(Simpson et al. 1989). Species that do not form seed banks will be solely dependent on
dispersal for the recolonisation of a site, which is generally a slow process (Luzuriaga et
al. 2005).

On a community level, vegetation types that accumulate seed banks recover quicker from
disturbances (Luzuriaga et al. 2005). However, not all the species in an existing
community are found in the seed bank (Crawford & Young 1998, Bossuyt et al. 2005,
Mengistu 2005) and species in the seed bank are not always found in the extant
vegetation. The composition of the seed bank determines whether the pre-disturbance
community is restored. If similarity is low between the seed bank and the aboveground
vegetation, there will be a change in composition of the community (Crawford &Young
1998, Baskin & Baskin 1998). It is therefore paramount that the seed bank of a
community is understood to inform any management or rehabilitation actions (Bossuyt et
al. 2005).
21
2.1.1 Classification of seed banks
Seed banks are classified according to the length of time they are able to survive in the
soil. Thompson & Grime (1979) described four types of soil seed banks found in
temperate regions. Species that have transient seed banks during summer and germinate
in autumn are Type I while species that have transient seed banks in winter and germinate
in the spring are Type II. Type III and type IV species have persistent seed banks, with
the difference being in the former type the majority of seeds germinate quickly with only
a small proportion entering into a persistent seed bank, as opposed to the latter group
where most seeds enter into a persistent seed bank and a minority germinate immediately.
As type III and IV have been found to be two ends of a continuum and little information
can be gained on longevity as the period of persistence may be anything from a few years
to many decades, a new classification was proposed.

Thompson et al. (1997) came up with a classification based solely on longevity. Species
with transient seed banks survive in the soil for less than a year, and no clear distinction
is made between summer- and winter-dormant types as this is irrelevant in tropical
environments. Species with persistent seed banks are divided into short-term and long-
term types, with seeds remaining viable for between one and five years and greater than
five years, respectively.

2.1.2 Composition and distribution of a seed bank
The size and density of a seed bank determines its importance to the functioning of a
plant community under which it occurs (Thompson 1992). Its composition and density is
determined by a number of factors, including fruit and flower production, seed rain
(Cumins & Miller 2002), seed dispersal, granivory (Hulme 1998, Auld & Denham 2001,
Barnes 2001), seed burial (Luzuriaga et al. 2005), germination and seed death (Simpson
et al. 1989). Management practises such as grazing (Kinloch & Friedel 2005a) and
burning (Cummins & Miller 2002) can also affect seed banks. The composition and
abundance of a seed bank is a reflection of what the community will look like after severe
disturbance (Luzuriaga et al. 2005).
22
The distribution of seed banks tend to be heterogeneous in landscapes (Crawford &
Young 1998, Page et al. 2006), with lower seed densities associated with clearings and
bare soil surfaces and higher seed densities in areas of perennial vegetation, depressions
and litter-covered soil surfaces (Guo 1998, Kinloch & Friedel 2005b). This is a result of
differential dispersal and granivory.

Seed dispersal increases the fitness of species and maintains populations by either
avoiding negative conditions underneath the parent plant, such as competition and
disease, or by exploiting beneficial microhabitats away from the parent plant (Simpson,
Leck & Parker 1989, Baskin & Baskin 1998, Wilson 1992). This allows species to
increase their geographical distribution, decreasing the risk of extinction. Dispersed seeds
may either become part of the seed bank, or may germinate immediately, in which case
many will perish before or after germination, as their depositional environment is either
environmentally unsuitable or they are targeted by herbivores (Stiles 1992). However,
some seeds will be dispersed to safe sites that are conducive to establishment.

The seeds of plants are dispersed in various ways, including autochory (self-dispersed),
anemochory (wind-dispersed), zoochory (animal-dispersed) and hydrochory (water-
dispersed) (van der Pijl 1972). Each of these dispersal mechanisms is accompanied by
morphological adaptions of the diaspore. Autochorous species forcibly eject their seeds
and have no other obvious adaptation to dispersal. Species which are anemochorous have
either wings, plumes or balloons to assist with transport by the wind, or have tiny seeds,
such as the Crassulaceae, Scrophulariaceae and Orchidaceae (van der Pijl 1972). Plants
can be dispersed by a variety of animals, including birds, mammals, reptiles and ants
(myrmechory). They can be ingested (endozoochory) or transported on the coats of
animals (exozoochory) (Fenner & Thompson 2005). Fruits that are dispersed by
endozoochory tend to be large, fleshy and generally attractive to animals. However, some
species with small seeds are also able to survive passage through the gut of an animal,
and are endozoochorous (Pakeman et al. 2002). Exozoochorous species have burrs,
spines or sticky substances that allow easy attachment to animals (Fenner & Thompson
23
2005). Seeds of myrmechorous species have edible eliosomes that act as a food source to
ants, causing them to be transported underground (Milewski 1982).

Seed banks can only contribute to the regeneration of a population and community if
there is a supply of safe sites that promote the emergence and establishment of new
individuals (Sigwela 2004). Directed dispersal allows seeds to be spread to microhabitats
that are beneficial to establishment.

2.1.3 Factors affecting the seed bank in Thicket
As Thicket is dominated by long-lived, stress-tolerant plant species (Cowling 1984) that
are able to reproduce vegetatively (Midgley & Cowling 1993), it can be assumed that the
main mechanism of regeneration would not be by a persistent seed bank, but by
vegetative expansion (Garwood 1989, Luzuriaga et al. 2005). This is especially true as
disturbance is of a generally low intensity in the natural functioning of Thicket, and does
not result in the complete removal of aboveground biomass (Kerley et al. 1995),
precluding the need for regeneration through a seed bank. Neither is there any evidence
of a persistent seedling bank (Grime & Hillier 1992).

Most woody species in Thicket produce fleshy fruits that are dispersed by birds or
mammals, compared to anemochorous species in the clearings or surrounding vegetation
(Pierce & Cowling 1991, Watson 2002). As most mammals and birds prefer the safety of
the dense cover of vegetation (Alcantara et al. 2000, Sigwela 2004), it would be expected
that the seed bank underneath the canopy of Thicket is different to the seed bank in
clearings. Underneath the canopy of Thicket, the seed bank should be dominated by
zoochorous species, as seeds with awns or wings should not be able to penetrate into the
bushclumps. The clearings would be expected to be dominated by anemochorous or
autochorous species (Watson 2002). The edge of the bushclumps should have the highest
species richness and density, as it is habitat for birds and mammals, as well as acting as a
barrier to wind-dispersed species.

24
The seeds of canopy Thicket species are quite large and are more likely to be consumed
by granivores (Fenner & Thompson 2005). However, many species produce large seed
crops (Cowling et al. 1997) and are mast seeders (Sigwela 2004), increasing the chance
of seeds becoming buried and entering the seed bank. Masting, as well as zoochory, are
traits selectively reinforced by granivory (Louda 1989). As most woody species are
absent from the seed bank, it is assumed that their seeds are not recalcitrant, and
germination has to occur immediately for successful establishment to occur (Sigwela
2004), similar to tropical forest trees (Garwood 1989). Individual plants need to produce
one surviving seedling per lifetime (Moles et al. 2003). Therefore for long-lived species,
low seed density in the seed bank is not a sign of poor persistence in the landscape.

However, most Thicket species expend a lot of energy flowering and fruiting regularly
and throughout the year (Bruton 1992, Watson 2002), with some benefit expected. Is it
simply to maintain the dispersers and other animals required by the community, or is
there a reward for that expenditure?

In Thicket, the microclimate underneath the canopy differs considerably from the
environment outside (Sigwela 2004). As most mammals in Thicket are browsers and
spend most of their time in the Thicket, zoochory would be the most beneficial
mechanism of dispersal in Thicket. Seeds will be directly dispersed into the microclimate
under the canopy of Thicket, preventing dispersal into clearings as both physical
conditions and soil structure is not beneficial to the establishment of seedlings there.
Thicket clump soils have higher organic (Hall 1997, Todkill 1997), moisture and nitrogen
content (Watson 2002) and lower light levels (Sigwela 2004), which will all promote the
successful emergence and establishment of seedlings.

Few seedlings of woody Thicket species have been recorded in open areas (La Cock
1992, Sigwela 2004). This is mainly due to the harsh environment. However, in more
mesic conditions where there is a more beneficial environment for seedling emergence
and establishment, are seeds dispersed into clearings? Or are there no seedlings because
of lack of dispersal? Thicket historically contained a large suite of possible dispersers,
25
ranging from the steenbok to the elephant, as well as a rich avifauna (Vlok et al. 2003).
Many of the larger mammals, and especially the megahebivores, have had their numbers
reduced or have been driven to extinction, or extirpated, from the landscape (Kamineth
2004).

Very few studies have been conducted on the seed banks of Thicket. In Xeric Succulent
Thicket, the seed bank is dominated by grasses and forbs (Sigwela 2004). This is similar
to what Pierce and Cowling (1991) found in Dune Thicket (Low & Rebelo 1998; Vlok et
al. 2003) in the southeastern Cape. In both types of thicket there was a low similarity
between the aboveground vegetation and the seed bank. The Thicket seed bank has a low
species richness and is relatively low seed density, and will therefore make a small
contribution to the regeneration and rehabilitation of the vegetation.

This chapter aims to further the understanding of the seed bank in the mesic Thicket
vegetation types that have not been studied previously. The effect of the dispersal
mechanism of a species on the composition of the seed bank underneath Thicket clumps
as opposed to openings is described.

The objectives of this study are to:
1. describe the growth forms and dispersal mechanisms of the germinable seed bank
of mesic Thickets;
2. describe the differences between the seed bank underneath the canopy, on the
edge of mesic Thicket clumps and in between clumps; and
3. determine the relationship between the seed bank and aboveground vegetation for
mesic Thicket types.

Hypotheses:
H1: Mesic Thicket seed banks are dominated by herbs and grasses;
H2: The seed bank underneath the canopy of mesic Thickets contains more woody
species, compared to interclump areas;
26
H3: The dominant dispersal mechanism of species found in the seed bank is zoochory
underneath the canopy, and anemochory in the interclump areas; and
H4: The number of seedlings of woody species will increase with an increase in rainfall
of a site.

2.2 Materials and Methods
2.2.1 Study sites
The initial selection of sampling sites for this study was based on the classification of
Thicket by Everard (1987) (Fig. 3). This classification was chosen because it was based
on characteristic species for each of the vegetation units, as well as being supported by
multivariate analysis. Although Everard (1987) identified four Thicket types, the Xeric
Succulent Thicket was excluded in this study as previous seed bank work had already
been done in this type (Sigwela 2004). For the remaining three vegetation types, three
study areas were selected in each. The Subtropical Thicket Ecosystem Planning (STEP)
Projects classification was also referred to, as it is more recent, and the classification that
best described the phytosociology of the sampling sites was used in the analysis of the
data (Fig. 4 and 5). Weather data for the closest weather stations to the sites was obtained
from Weather SA (2008).

Mesic Kaffrarian Thicket
Umtiza Nature Reserve (MKT1), Fort Jackson (MKT2) and Sittingbourne (MKT3) are
situated near to each other in the East London area. The region has a warm and moist
climate. Soils are derived from the dolerites intrusive in the Beaufort group (Everard
1987). Umtiza is in a declared provincial nature reserve, but due to the dense nature of
the vegetation, the only clearings were along a relatively busy road, near to urban areas.
Fort Jackson is located in a commercial farming area, whereas Sittingbourne is
communal grazing land. All three sites fall within the Buffels Thicket (BT) of STEP
(Vlok et al. 2003), and there codes are BT1, BT2 and BT3 respectively. Rainfall for East
London is 921 mm per annum (Weather SA 2008).

27
Mesic Kaffrarian Thicket
Mesic Succulent Thicket
Xeric Kaffrarian Thicket
Xeric Succulent Thicket
0 50 100 25 km

Figure 3 Geographical location of study sites according to Everards (1987) classification of Thicket, and climate data of the nearest weather stations.

J F M A M J J A S O N D
0
40
80
120
160
200
R
a
in
f
a
ll (
m
m
)
0
4
8
12
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28
T
e
m
p
e
r
a
t
u
r
e
(
C
)

0
40
80
120
160
200
R
a
in
fa
ll (m
m
)
0
4
8
12
16
20
24
28
32
T
e
m
p
e
r
a
tu
r
e
(
C
)

0
40
80
120
160
200
R
a
in
f
a
ll (
m
m
)
0
4
8
12
16
20
24
28
T
e
m
p
e
r
a
t
u
r
e
(
C
)

0
40
80
120
160
200
R
a
in
f
a
ll (
m
m
)
0
4
8
12
16
20
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28
T
e
m
p
e
r
a
t
u
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(
C
)
28
")")
") ") ")
")")")")")
")")")")")
^_
^_^_^_^_
!(!(
!( !(!(
!(!(!( !(!(
^_^_^_^_^_
#* #*#*#*#*
#*#*#*
#*#*
Sites
") Umtiza (BT1)
") Sittingbourne (BT2)
") Fort Jackson (BT3)
^_
Gamtoos (GT1)
^_
Loerie (GT2)
#* Shamwari (AT2)
#* Thomas Baines (AT2)
!( Welbedachtsfontein (ST1)
!( Wells Estate (ST2)
STEP thicket type
DUNE THICKET
ARID THICKET
THICKET
VALLEY THICKET
0 50 100 25 km

Figure 4 Geographical location of study sites according to STEPs (Vlok et al. 2003) structural
vegetation units of Thicket.
")")
") ") ")
")")")")")
")")")")")
^_
^_^_^_^_
!(!(
!( !(!(
!(!(!( !(!(
^_^_^_^_^_
#* #*#*#*#*
#*#*#*
#*#*
Sites
") Umtiza (BT1)
") Sittingbourne (BT2)
") Fort Jackson (BT3)
^_
Gamtoos (GT1)
^_
Loerie (GT2)
#*
Shamwari (AT2)
#*
Thomas Baines (AT2)
!( Welbedachtsfontein (ST1)
!( Wells Estate (ST2)
STEP thicket type
ALBANY
ALGOA
BUFFELS
CENTRAL
EASTERN
ESCARPMENT
FAREASTERN
FISH
GAMTOOS
GROOT
KEI
SUNDAYS
TRANSFISH
0 50 100 25 km

Figure 5 Geographical location of study sites according to STEPs (Vlok et al. 2003) biogeographical
units ofThicket.
29
Xeric Kaffrarian Thicket
Gamtoos (XKT1) is situated between Port Elizabeth and Jeffreys Bay. As it is near the
coast rainfall is relatively high (Cowling 1984). Soils consist of recently consolidated
calcareous dune sands of the Nanaga formation (GSSA 1991). Gamtoos is located in
Gamtoos Thicket, and its code according to STEP is GT1.

Thomas Baines Nature Reserve (XKT2) is a provincial nature reserve located south of
Grahamstown. Sandy soils are derived from quartzites of the Witpoort formation (Vlok
& Euston-Brown 2002). Shamwari Game Reserve (XKT3) is a private game reserve
situated north-east of Port Elizabeth. The southern part of the reserve, where the study
sites were located, is dominated by the Sundays River formation resulting in shallow soils
underlain by calcrete (OBrien 2004). Both sites experience a warm climate with bimodal
rainfall, with maxima occurring in September to October and February to March. Mean
annual rainfall is 420 mm in Shamwari (OBrien 2004). The nearest weather station to
Thomas Baines is Grahamstown, where mean annual rainfall is 505 mm (Weather SA
2008), although the reserve will have lower rainfall as it is situated in a drier area.
Thomas Baines and Shamwari occur in Albany Thicket (AT), and their site codes are
AT1 and AT2, respectively.

Mesic Succulent Thicket
The PPC Loerie limestone quarry (MST1) is situated within the Gamtoos River Valley,
near the town of Loerie. It has a warm, fairly dry climate (Hall 2000). Soils are derived
from limestone and phyllites of the Cango formation, and are deep with uniformly red,
apedal subsoils of the Hutton formation with slightly acidic to neutral topsoils (Cowling
1984, Hall 2000). Loerie is situated within Gamtoos Thicket, and GT2 is its site code.
Rainfall at the nearest weather station in nearby Hankey is 440 mm.a
-1
(Weather SA
2008).

Welbedachtsfontein (MST2) and Wells Estate (MST3) are situated close to each other,
north of Port Elizabeth. Welbedachtsfontein is a farm is situated along the Addo Road,
bordering the Grassridge. Wells Estate is situated within 100 m from the coast, between
30
the city of Port Elizabeth and the new Ngqura harbour. Mean annual rainfall is 438 mm at
the harbour, decreasing to 417 mm at Uitenhage, situated further inland (Weather SA
2008). Soils are loamy or clayey, predominantly derived from the Bluewater Bay
formation, although Welbedachtsfontein is more complex, including the Kirkwood,
Alexandria, Salnova and Sundays River formations (GSSA 1991). Since sampling for
this study took place, Wells Estate has been cleared for the Coega Industrial
Development Zone. Both locations are situated in Sundays Thicket, and their site codes
are ST1 and ST2 respectively.

2.2.2 Field Sampling
At each study site, transects (n = 5) were used to determine species composition and
cover. The location of each transect was positioned at the edge of a Thicket clump,
allowing the identification of the effect of the microclimate on the seed bank of Thicket.
Each transect consisted of three equidistant 25 x 25 cm quadrats. The first quadrat at the
beginning of the transect was placed in the interclump area (OP), with the next one
positioned at the edge of the Thicket clump (ED). The location of the third quadrat below
the Thicket canopy (CA) was positioned at equal distance from the quadrat at the edge as
the OP and ED quadrats. The cover abundance of all plants was determined both for the
Thicket clump and interclump area. At each quadrat, litter depth was measured, and soil
and air temperature measured using a Bailey thermocouple (Sensortek, BAT-12). Light
was measured with terrestrial light meter (Licor, Li-189). Seedlings found along the
transect were counted and uprooted for later identification. Uprooting was required for
the identification of whether the propagules were a seedling or ramet. Seedlings of
grasses and weedy annual species were excluded from the analysis, as identification of
their seedlings was deemed imprecise. Where seedlings could not be identified down to
species level, they were grouped according to their genus.

Soil samples were collected by excavating each quadrat to a depth of 15 cm. They were
transported in plastic bags back to the laboratory, where they were stored until July,
2006. The bags were stored open to prevent anoxic conditions due to bacterial activity.
31

Voucher specimens of the plants recorded are located at the Ria Olivier Herbarium in the
Botany Department of Nelson Mandela Metropolitan University. The Mesic Succulent
Thicket sites were sampled in April, the Xeric Kaffrarian Thicket sites in May, and the
Mesic Kaffrarian Thicket sites in June, 2006.

2.2.3 Laboratory study
The soil was returned to the laboratory where it was well mixed and weighed. Half of the
sample was placed in seedling trays in an enclosed glasshouse in July, 2006. The trays
were rotated on a monthly basis. Two control trays consisting of autoclaved potting soil
were randomly placed in between each sites trays to account for seed contamination.
Tarchonanthus camphoratus, Brachylaena discolor, Conyza albida, C. bonariensis and
Oxalis corniculata came up in the control trays, and were excluded from the results. All
seed trays were checked weekly. Emergent seedlings of herbs and grasses were left until
identification was possible, counted and then removed by uprooting. Seedlings of shrubs
and succulents were transplanted into pots and kept for later identification. Once
germination stopped or greatly diminished, all plant material was removed and the soil
was mixed to bring buried seeds to the surface. The experiment continued for 18 months,
until December 2007.

The species in the seed bank were classified according to growth form as follows: woody
shrub or tree (woody), herb, succulent, grasses and sedges (graminoid), bulb (geophyte),
or climber. Fruit morphology (dehiscent, indehiscent or fleshy) and dispersal mechanism
(anemochorous, autochorous or zoochorous) was taken from available literature
(Thiselton Dyer 1900, Pooley 1998, Van der Planck 1998, 1999, Van Oudtshoorn 1999,
Leistner 2000, Manning et al. 2001 Coates Palgrave 2002).

Species richness (d) was determined using Margalefs index (d=[S-1]/log[N]), for both
aboveground vegetation and the seed bank, where S is the total number of species at a
site and N is the total number of species observed at all sites. The Shannon-Wiener
32
function (H) was used as a measure of diversity, and Simpsons index (C) as an
estimation of species dominance. Srensen's (1948) coefficient of community (CC) was
used to test for the similarity between the seed bank and aboveground vegetation at each
corresponding site, where CC = 2 S
S
/(S
a
+S
b
) in which S
s
is the number of species shared
by two samples, S
a
is the number in the first and S
b
the number in the second sample.

Detrended Correspondence Analysis (DCA) was used (CANOCO, Ter Braak 1987) to
identify the relationships between the vegetation of the different sampling sites, as well as
the seed banks of the three microenvironments at each sampling site. The four replicates
were combined for the seed bank samples. Factorial ANOVA with the Tukey post-hoc
test, or Kruskal-Wallis analysis by ranks (for non-parametric data) was used to test for
significance between the environmental variables and diversity indices of both the seed
bank and aboveground vegetation, using Statistica 8 (Statsoft 2007).

2.3 Results
2.3.1 Aboveground Vegetation
Ordination of the vegetation samples revealed the superior explaining power of the STEP
(Vlok et al. 2003) classification compared to Everard (1987), although certain overlap
and separation of vegetation types still occurred. It was decided to use STEP for the rest
of the chapter.

The canopy of Buffels Thicket (BT) was dominated by Acacia karroo, Grewia
occidentalis, Scutia myrtina, and Diospyros dichrophylla (Table 2). Other species found
at all three study sites were Abutilon sonneratianum, Burchellia bubalina, Eragrostis sp.,
Hippobromus pauciflorus, Cheilanthes viridis, Rhoicissus tridentata, Rhus chirindensis
and Tecoma capensis.

The dominant species in Albany Thicket (AT) were Euclea undulata, Schotia afra,
Ptaeroxylon obliquum and Azima tetracantha (Table 2). Species shared between both
33
Table 2 Mean (n = 5) cover of species in aboveground vegetation with greater than 1% cover. BT =
Buffels Thicket; AT = Albany Thicket; ST = Sundays Thicket; GT = Gamtoos Thicket.
Species BT1 BT2 BT3 AT1 AT2 ST1 ST2 GT1 GT2
Acacia karroo Hayne 16.0 8.0 17.0 1.4 1.0 2.1
Acokanthera oppositifolia (Lam.) Codd 2.0
Aloe africana Mill. 1.4 * * * *
Aloe ferox Mill. 2.1
Aloe pluridens Haw. * 1.8 *
Azima tetracantha Lam. 4.0 7.4 * * 1.1 1.8
Brachylaena elliptica (Thunb.) DC. 2.0
Brachylaena ilicifolia (Lam.) Phill. &
Schweick 3.0 6.0 1.0 2.0
Burchellia bubalina (L.f.) Sims * 2.1 2.0
Capparis sepiaria L. var. citrifolia (Lam.)
Tlken 5.4 * * 2.4 * 1.4
Carissa bispinosa (L.) Desf. ex Brenan * * 1.5 0.9 * 1.0 *
Chrysanthemoides monilifera (L.)
T.Norl. subsp. monilifera 3.0
Clausena anisata (Willd.) Hook.F. ex
Benth. * 3.0
Crassula mesembryanthoides (Haw.)
D.Dietr. * 1.0 *
Crassula ovata (Mill.) Druce * * * 2.4
Crassula perforata Thunb. 0.1 * 1.0
Diospyros dichrophylla (Gand.) De
Winter 11.0 50.0 5.1 *
Ehretia rigida (Thunb.) Druce subsp.
Rigida 10.0 9.0 4.4 2.0 2.1 1.4
Euclea undulata Thunb. 10.4 13.0 14.0 30.0 14.0 2.4
Euphorbia ledienii A.Berger 4.8 1.0
Grewia occidentalis L. * * 10.0 3.4 1.1
Grewia robusta Burch. 3.0 * 9.2
Gymnosporia capitata (E.Mey. ex
Sond.) Loes. 6.0 4.0 16.0 * 19.0
Gymnosporia heterophylla (Eckl. &
Zeyh.) Loes. 1.2 7.4 2.1
Helichrysum teretifolium (L.) D.Don 5.0 3.1 *
Hippobromus pauciflorus (L.f.) Radlk. 16.0 * * * 1.0 18.4
Hypoestes aristata (Vahl) Sol. ex Roem.
& Schult. 4.0 *
Hypoestes forskaolii (Vahl) R.Br. 19.0 2.1 * 6.0 9.6 12.0
Jasminum angulare Vahl 1.0
Lauridia tetragona (L.f.) R.H.Archer 4.0
Maerua cafra (DC.) Pax * 1.0
Aptenia cordifolia (L.f.) Schwantes * 2.1
Mystroxylon aethiopicum (Thunb.) Loes.
subsp. aethiopicum * * 1.0 2.0
Olea europaea L. subsp. africana (Mill.)
P.S.Green * 2.0 * 7.4
Opuntia ficus-indica (L.) Mill. * * * * 1.0
Panicum deustum Thunb. * * * * * 1.4 *
Pappea capensis Eckl. & Zeyh. 3.0 3.2 * *
Plumbago auriculata Lam. * 2.5 1.0 1.8
Polygala myrtifolia L. 1.0
Portulacaria afra Jacq. 1.0 *
Ptaeroxylon obliquum (Thunb.) Radlk. 4 9 11 *

cont.
34
Species BT1 BT2 BT3 AT1 AT2 ST1 ST2 GT1 GT2

Pterocelastrus tricuspidatus (Lam.)
Walp.

*

1.0
Putterlickia pyracantha (L.) Szyszyl. 1.0 * 2.1 3.0
Rhoiacarpos capensis (Harv.) A.DC. * * 2.1 * 1.0
Rhoicissus tridentata (L.f.) Wild &
R.B.Drumm. subsp. tridentata * * * 1.4 * * * 6.0 5.4
Rhus chirindensis Baker f. 6.0 * 1.0
Rhus dentata Thunb. 8.0 *
Rhus laevigata L.
Rhus pallens Eckl. & Zeyh. 6.0 1.0 * * 3.1 4.4
Rhus pterota C.Presl 2.0 6.0 4.0 5.0 2.0 2.0
Rhus tomentosa L. * * * 8.0
Schotia afra (L.) Thunb. var. afra * 10.0 6.0 18.4 11.0 *
Scolopia mundii (Eckl. & Zeyh.) Warb. * 11.4
Scutia myrtina (Burm.f.) Kurz 12.0 26.0 32.0 4.1 * 14.0 23.0
Sideroxylon inerme L. subsp. inerme * 2.0 2.0 8.0 17.0 10.4
Tecoma capensis (Thunb.) Lindl. 12.0 * *
Zanthoxylum capense (Thunb.) Harv. 10.0
Ziziphus mucronata Willd. 4.0 10.0
Trimeria grandifolia (Hochst.) Warb. 14.0
Rapanea melanophloeos (L.) Mez 15.0
Zygophyllum morgsana L. * 2.2
* present at location with less than 1% cover.

sites are Aloe africana, Aptenia cordifolia, Capparis sepiaria, Carissa bispinosa,
Commelina africana, Crassula mesembryanthemoides, Gymnosporia heterophylla,
Opuntia ficus-indica, Panicum deustum, Rhoicissus tridentata, Rhus pterota, Sansevieria
hyacinthoides, Sarcostemma viminale subsp. viminale and Sideroxylon inerme.

Euclea undulata, Pappea capensis, Schotia afra, Rhus pterota and Sideroxylon inerme
were the dominant species in Sundays Thicket (ST) (Table 2). Shared species between
the two sampling sites included Aloe pluridens, Asparagus africanus, A. racemosus, A.
suaveolens, A. subulatus, Brachylaena ilicifolia, Azima tetracantha, Capparis sepiaria,
Carissa bispinosa, Crassula cotyledonis and C. tetragona, Euphorbia ledienii,
Fockea edulis, Gymnosporia capitata, Hypoestes forskaolii, Kedostris nana, Mystroxylon
aethiopicum subsp. aethiopicum, Panicum deustum, Putterlickia pyracantha,
Rhoiacarpus capensis, Rhus pallens, Sarcostemma viminale, and Viscum rotundifolium.

35
The canopy of Gamtoos Thicket (GT) was dominated by Euclea undulata, Hippobromus
pauciflorus, Scutia myrtina and Sideroxylon inerme (Table 2). Acacia karroo, Aloe
africana, Azima tetracantha, Capparis sepiaria, Carissa bispinosa, Ehretia rigida,
Hypoestes forskaolii, Olea europaea subsp. africana, Opuntia ficus-indica, Panicum
deustem, Pappea capensis, Plumbago auriculata, Putterlickia pyracantha, Rhoiacarpos
capensis, Rhoicissus tridentata, Rhus pallens and Rhus pterota were some species shared
between both sites. Species that were dominant at Loerie but absent or not dominant at
Gamtoos were Schotia afra and Scolopia mundii, whereas the reverse was true for
Grewia robusta and Gymnosporia capitata.

Only one species was recorded at all sampling sites, namely Rhoicissus tridentata.
Species found in at least one sampling site of each of the four vegetation types were
Asparagus africanus, Carissa bispinosa, Cussonia spicata, Ehretia rigida, Hermannia
altheoides, Hippobromus pauciflorus, Hypoestes forskaolii, Panicum deustum, Rhus
pallens and Scutia myrtina.

Ordination of samples indicates that Mesic Kaffrarian Thicket differs substantially from
Xeric Kaffrarian and Mesic Succulent Thicket, separating from the latter in the major
ordination axis (x-axis; Fig. 6). The three Mesic Kaffrarian Thicket sites separate from
each other on the y-axis. One transect in Fort Jackson (MKT3), however, was more
similar to Sittingbourne (MKT2), as it shared Diospyros dichrophylla as a dominant
canopy species with over 50% cover. Umtiza (MKT1) separates into two groups, which
were a distance from each other, with the two samples near the origin containing more
forest elements such as Rapanea melanophloeos. The Xeric Kaffrarian Thicket and Mesic
Succulent Thicket have different phytosociology, except for samples of Loerie (MST1)
and Gamtoos (XKT1), both of which showed more similarity to the other vegetation type

When STEPs classification (Vlok et al. 2003) is used, the locations sort out according to
the biogeographical regions. Gamtoos (GT1) and Loerie (GT2) form their own group,
separate from the other sampling sites of Xeric Kaffrarian Thicket and Mesic Succulent
Thicket, respectively (Fig. 7). There is a close phytosociological relationship between
36
-1 6
-
1
4
MKT1 MKT2 MKT3
MST1 MST2 MST3
XKT1 XKT2 XKT3

Figure 6 Detrended Correspodence Analysis (DCA) of study sites, using coding according to
Everard's (1987) classification. MKT = Mesic Kaffrarian Thicket; XKS = Xeric Kaffrarian Thicket;
MST = Mesic Succulent Thicket. (x-axis eigenvalue = 0.745, y-axis eigenvalue = 0.404. Cumulative
percentage variance = 19.8%, Sum of all eigenvalues = 5.813)

-1 6
-
1
4
BT1_T BT2_T BT3_T GT1_T
GT2_VT ST1_VT ST2_VT AT1_T
AT1_VT AT2_VT

Figure 7 Detrended Correspodence Analysis (DCA) of study sites, using coding according to STEP's
biogeographic classification (Vlok et al. 2003). AT = Albany Thicket; BT = Buffels Thicket; GT =
Gamtoos Thicket; ST = Sundays Thicket. Sites are also coded for Thicket (T) and Valley Thicket
(VT). DCA statistics as in Figure 6.

Gamtoos
Sundays
Albany
Buffels
37
Gamtoos Thicket and Sundays Thicket, with some overlap between them. Albany
Thicket has three outliers, as Acacia karroo, Brachylaena ilicifolia, Grewia occidentalis
and Olea europea subsp. africana had greater cover at the three transects that fall with
Albany Valley Thicket, compared to Albany Thicket. The sites do not sort out according
to the structural types of Thicket, as Thicket (T) units of Gamtoos (GT1) and Thomas
Baines (AT1) are found within the rest of the Valley Thicket (VT) samples.

As sampling occurred at different times for the various vegetation types, it is not possible
to compare their environmental parameters. However, comparison may be made between
the three microhabitats at each study site. All environmental measures differed
significantly between microhabitats for the four vegetation types (Table 3). The
microhabitat beneath the canopy was significantly cooler than the open, both in terms of
the soil and the air, as well as having much reduced light, as little as 1-2% of the opening
(Fig. 8). The litter layer was significantly deeper below the canopy compared to the open.
The edge may significantly differ from both of the other microhabitats, but always had an
intermediate value for all four environmental parameters

A total of 63 species were found as seedlings (Table 4). Most species consisted of only a
few individuals limited to one or two sites. The most widespread and numerous species
was the herb Hypoestes sp. Fort Jackson (BT3) had both the greatest species richness and
seedling abundance, and Sundays Thicket (ST) the least. There was a large difference
between the two study sites of both Albany Thicket (AT) and Gamtoos Thicket (GT),
with both Gamtoos (GT1) and Thomas Baines (AT1) had much greater species richness
and seedling abundance than Loerie (GT2) and Shamwari (AT2). The majority of
seedlings were of woody species, followed by succulents (Fig. 9). Both the total number
and relative proportion of woody seedlings decreased from east to west (higher rainfall to
lower rainfall), with a concurrent increase in the number of succulent seedlings.

There was an increase in species richness (d) and diversity (H) from east to west, as
Buffels Thicket had the lowest and Sundays Thicket had the highest richness and
Gamtoos Thicket was the most diverse vegetation type (Table 5). There was a large
38
Table 3 F-values of environmental measures between the three microhabitats of the four Thicket
types. AT = Albany Thicket; BT = Buffels Thicket; GT = Gamtoos Thicket; ST = Sundays Thicket.
Site Enviro measure F df p
BT air temp 7.44 2 0.002
soil temp 22.05 2 <0.001
light 104.02 2 <0.001
litter 3.29 2 0.047

AT air temp 12.37 2 <0.001
soil temp 209.36 2 <0.001
light 200.51 2 <0.001
litter 3.64 2 0.04

ST air temp 21.26 2 <0.001
soil temp 131.44 2 <0.001
light 274.11 2 <0.001
litter 15.99 2 <0.001

GT air temp 5.6 2 0.009
soil temp 25.69 2 <0.001
light 446.73 2 <0.001
litter 8.73 2 0.001

difference between the diversity indices of Albany Thicket, as Thomas Baines (AT2) had
much higher species richness and diversity compared to Shamwari (AT2). Species
dominance (C) was almost double in Albany Thicket and Gamtoos Thicket, compared to
the other two vegetation types.
39
BT AT ST GT
0
10
20
30
40
A
i
r

t
e
m
p
e
r
a
t
u
r
e

(
C
)
Opening
Edge
Canopy
BT AT ST GT
0
10
20
30
40
50
S
o
i
l

t
e
m
p
e
r
a
t
u
r
e

(
C
)
Opening
Edge
Canopy

a) b)
BT AT ST GT
0
400
800
1200
1600
2000
L
i
g
h
t

(
m
o
l
.
m
-
2
.
s
-
1
)
Opening
Edge
Canopy
BT AT ST GT
0
1
2
3
4
5
L
i
t
t
e
r

d
e
p
t
h

(
c
m
)
Opening
Edge
Canopy

c) d)
Figure 8 Air temperature (a), soil temperatures (b), light (c) and litter depth (d) in three
environments (canopy, edge and opening) of study sites. AT = Albany Thicket; BT = Buffels Thicket;
GT = Gamtoos Thicket; ST = Sundays Thicket.

a
ab
b
b
bc
a
a
c
c
a ab
b
a
bc
c
a
bc
c
a
bc
c
a
ab
b
a
bc
c
a
b
c
a
b
c
a
b
c
a
ab
b
a
b
b
a
b
c
a
ab
b
40
Table 4 The number and growth form (G) of seedlings collected along transects (n=5) at each study
site and no. of sites (no.) where seedlings of each species occurs. AT = Albany Thicket; BT = Buffels
Thicket; GT = Gamtoos Thicket; ST = Sundays Thicket. (Wo = woody, He = herb, S = succulent, Gr
= graminoid, Cl = climber, Ge = geophyte)
Seedlings G
BT
1
BT
2
BT
3
GT
1
AT
1
AT
2
GT
2
ST
1
ST
2
n
o.
Abutilon sonneratianum (Cav.) Sweet He 2 1 2
Acacia karroo Hayne Wo 2 1
Acalypha villicaulis Hochst. He 1 1
Aloe africana Mill. S 1 1
Aloe ferox Mill. S 1 2 2
Aloe pluridens Mill. S 4 1 2
Aptenia cordifolia (L.f.) Schwantes S 1 1
Asparagus sp. Wo 1 1
Azima tetracantha Lam. Wo 2 1 2
Burchellia bubalina (L.f.) Simms Wo 4 2 2
Canthium sp. Wo 1 1
Cassine peragua L. Wo 1 1 2
Celtis africana L. Wo 5 1
Commelina sp. Wo 2 7 2
Crassula expansa Dryand. S 1 1 1 3
Crassula ovata (Mill.) Druce S 1 1 2
Crassula perforata Thunb. S 1 1
Crassula spathulata Thunb. S 1 1
Cussonia sp. Wo 3 1 1 3
Diospyros dichrophylla (Gand.) De Winter Wo 1 2 2
Dovyalis rhamnoides (Burch. Ex DC.) Burch. &
Harv.
Wo 1 7 2
Ehretia rigida Thunb. Druce Wo 3 4 2
Euclea sp. Wo 1 1 1 3
Euphorbia ledienii A. Burger. S 1 1
Euphorbia triangularis Desf. S 11 1
Galenia pubescens(Eckl. & Zeyh.) Druce S 1 1
Galopina circaeoides Thunb. He 4 1 2
Gasteria bicolor Haw. var. bicolor S 1 2 2 3
Grewia occidentalis L. Wo 2 3 3 2 4
Gymnosporia sp. Wo 13 1
Helichrysum teretifolium (L.) D. Don Wo 2 1 2
Hippobromus pauciflorus (L.f.) Radlk. Wo 1 2 2
Hypoestes sp. He 15 11 2 7 2 2 6
Hypoxis stellipilis Ker Gawl. Ge 1 1
Jasminum angulare Vahl Wo 2 2 2
Kalanchoe rotundifolia (Haw.) Haw. S 1 1
Kedostris nana (Lam.) Cogn. Cl 2 3 2
Ledebouria revoluta (L.f.) Jessop Ge 1 1
Mystroxylon aethiopicum (Thunb.) Loes. Subsp.
aethiopicum
Wo 1 1
Ornithogalum tenuifolium F. Delaroche. subsp.
tenuifolium
Ge 1 1

Pelargonium peltatum ( L.) LHerit
He 3 2 2
Plectranthus madagascariensis (Pers.) Benth. var.
madagascariensis
He 7 3 2
Plumbago auriculata Lam. Wo 4 1
cont.
41
Seedlings G
BT
1
BT
2
BT
3
GT
1
AT
1
AT
2
GT
2
ST
1
ST
2
n
o.

Pseudognaphalium luteo-album (L.) Hilliard & B.R.
Burtt
He 2 2 2
Putterlickia pyracantha (L.) Szyszyl. Wo 2 1
Rhus sp. Wo 3 3 5 1 4
Sansevieria aethiopica (L.) Druce Ge 1 1 2
Sarcostemma viminale (L.) R.Br. subsp. viminale S 1 1 2
Schotia afra (L.) Thunb. var. afra Wo 1 1
Sebaea macrophylla Gilg He 1 1
seedling1 Wo 1 1
seedling2 Wo 1 1
seedling3 Wo 1 1
seedling4 S 1 1
seedling5 Wo 1 1
seedling6 He 1 1
Senecio pterophorus DC. He 2 1
Senecio radicans(L.f.) Sch.Bip. S 1 1
Sideroxylon inerme L. subsp. inerme Wo 4 1
Solanum sp. Wo 2 7 2
Tecoma capensis (Thunb.) Lindl. Wo 1 1
Vepris lanceolata Comm. ex A.Juss Wo 2 1
Ziziphus mucronata Willd. Wo 2 1 2
species richness 11 15 24 14 17 5 8 5 6
total no. of seedlings 33 40 66 37 37 7 14 9 9
no. of seedlings m
-2
2.4 4.0 8.2 4.4 0.8 1.3 1.7 1.9 4.5

2.3.2 Composition of the seed bank
A total of 114 species emerged from the seed bank, with most of them having a low mean
number of individuals (Table 6). Buffels Thicket had both the greatest species richness
and species diversity, compared to the other Thicket types that had similar values. The
seed bank in Buffels Thicket was dominated by Centella asiatica, Digitaria sanguinalis,
Gamochaeta pennsylvanica, Helichrysum cymosum, Panicum maximum and Senecio
polyanthemoides. Umtiza (BT1) had high abundances of Cyphia sylvatica, Priva meyeri
var. meyeri and Sporobolus africanus. Spermacoce natalensis, Pseudognaphalium luteo-
album and Cyphia sylvatica were dominant species in Sittingbourne (BT2), wheras
Galopina circaeoides dominated at Fort Jackson (BT3).

42
BT AT ST GT
0
20
40
60
80
100
P
r
o
p
o
r
t
i
o
n

g
r
o
w
t
h

f
o
r
m
woody
succulent
herb
climber
geophyte

Figure 9 The relative growth form composition of seedlings found at study sites. AT = Albany
Thicket; BT = Buffels Thicket; GT = Gamtoos Thicket; ST = Sundays Thicket.

Table 5 Species richness (d), species diversity (H) and species dominance (C) of the aboveground
vegetation of sampling sites.
Site d H' C
Buffels Thicket 1 13.34 2.57 11.91
mean 11.12 2.07 6.97
Albany Thicket 1 29.15 2.99 13.15
Albany Thicket 2 19.58 2.26 13.43
mean 24.36 2.63 13.29
Sundays Thicket 1 22.29 2.47 7.74
Sundays Thicket 2 28.98 2.45 6.07
mean 25.63 2.46 6.90
Gamtoos Thicket 1 21.40 2.92 13.15
Gamtoos Thicket 2 25.41 2.90 12.71
mean 23.41 2.91 12.93
43
Crassula expansa, Panicum maximum and P. deustum dominate the seed bank of Albany
Thicket (AT). Nemesia affinis was also dominant at Shamwari (AT2) and Sutera
polyantha co-dominant at Thomas Baines (AT1). The former four species are dominant
in the seed bank of Sundays Thicket (ST) together with Cynodon dactylon, as well as
Nemesia affinis dominating Welbedachtsfontein (ST2).

Dominant species in the germinable seed bank of Gamtoos Thicket (GT) are P.
maximum, C. dactylon, Ehrharta erecta, Mariscus congestus and Polycarpa
tetraphyllum. Galopina circaeoides and Digitaria natalensis are important species in the
germinable seed bank of Loerie (GT2) that are absent from Gamtoos (GT1).

Seed density (F = 7.93, d.f. = 3, p < 0.001) and species richness (F = 12.54, d.f. = 3, p <
0.001) differed significantly between sampling sites, but not between the different
microsites (Table 7). Albany, Sundays and Gamtoos Thicket differed from that of Buffels
Thicket (p = 0.001). Ordination of the germinable seed bank of the various vegetation
units shows a similar phytosociological relationship as that of the aboveground
vegetation, although there is greater similarity between Albany and Sundays Thicket
(Fig. 10). There is a clear separation along the major x-axis between Buffels Thicket and
the other Thicket units, with the various Buffels Thicket types separating along the y-
axis. The largest differences between the three environments occur in Gamtoos Thicket.

The seed bank of all Thicket types were dominated by herbs and grasses (Fig. 11).
Woody species were more common in Buffels Thicket while succulent species become
more important in the other three vegetation units, except under the canopy of Gamtoos
Thicket, where succulents were absent and woody species co-dominant. Geophytes and
climbers were rare. The edge of the Thicket in Loerie (MST1) has the highest number of
geophytes compared to all sites, containing four species, three species of Oxalis (O.
caprina, O. incarnata and O.punctata) and Ornithogalum tenuifolium subsp. tenuifolium.
Woody species made their largest contribution to the seed bank underneath the canopy in
Buffels and Gamtoos Thicket, with the reverse, largest contribution in the open, true in
Albany and Sundays Thicket. No clear pattern emerged for succulents.
44
Table 6 Mean (n = 4) number of seeds per m
2
, Growth form (G) and dispersal mechanism (DM) of the dominant species in the germinable seed bank of
the three microsites of the sampling sites. (CA = canopy, ED = edge, OP = open). (v = present in aboveground vegetation, sb = present in seed bank).
Small letters indicate significant differences between vegetation types for seed abundance and species richness.
G DM Buffels Thicket Albany Thicket Sundays Thicket Gamtoos Thicket
CA ED OP CA ED OP CA ED OP CA ED OP
Cyphia sylvatica Eckl. Cl W 47 195 64 v
Digitaria sanguinalis (L.) Scop. Gr W 48 131 144 v sb
Priva meyeri Jaub. & Spach var meyeri He A, Z 64 125 0
Sporobolus africanus (Poir.) Robyns & Tournay Gr A 27 64 89 v sb sb
Pseudognaphalium luteo-album (L.) Hilliard & B.R. Burtt He W, Z 16 64 35 sb sb
Galopina circaeoides Thunb. He A 5 21 64
Spermacoce natalensis Hochst. He A 32 45 24
Senecio inaequidens DC. He W 19 40 32 sb sb
Centella asiatica (L.) Urb. He Z 13 11 31
Selago corymbosa L. Wo W 11 24 5
Crassula expansa Dryand. S W 14 38 220 24 0 20 v sb
Panicum maximum Jacq. Gr W, Z sb 120 114 84 v 39 54 92 4 32 108
Nemesia affinis Benth. He W 84 8 4 208 192 18
Panicum deustum Thunb. Gr W, Z v 24 48 20 v 4 28 2 v v sb
Sutera polyantha (Benth.) Kuntze He A sb 0 28 6 sb sb
Cynodon dactylon (L.) Pers. Gr W v sb sb 48 24 20 10 36 56
Polycarpon tetraphyllum (L.) L. He W sb 0 4 84
Ehrharta erecta Lam. var. natalensis Stapf Gr W sb 14 12 48
Mariscus congestus Vahl Gr W, Z sb 38 12 12
Total no. of seeds m
416 1055 733 a 378 350 378 b 383 352 236 b 130 242 476 b
Total no. of species
39 46 38 a 21 27 12 b 16 12 18 b 16 20 26 b
45

-1.0 6.0
-
0
.
5
3
.
5

Figure 10 Detrended Correspondence Analysis (DCA) of the germinable seed bank of the three
microsites of the study sites. Blues are Buffels Thicket, Reds are Albany Thicket, Yellows are
Sundays Thicket and Purples are Gamtoos Thicket. (Filled circles = canopy, Hatched circles = edge,
Empty circles = open).

Anemochory was the dominant mechanism of dispersal at all three vegetation types, and
in all microenvironments, with zoochory co-dominant except in the open samples of
Sundays and Gamtoos Thicket (Fig. 12). Autochory was the least common. There was a
decrease in the relative abundance of zoochorous species from the canopy to the open.

The species richness and diversity in the germinable seed bank was higher in Buffels
Thicket compared to the other types of Thicket (Table 8). Highest species richness was
recorded at the edge of Thicket for Buffels and Albany Thicket, but in the open for
Sundays and Gamtoos Thicket. No clear pattern was identified for species diversity.

Table 7 F-values for differences in seed density and species richness of the germinable seed bank
between the four vegetation types and three microhabitats.
Seed density Species richness
F-value df p F-value df p
site 7.93 3 <0.001 12.54 3 <0.001
microhabitat 2.80 2 ns 1.75 2 ns
46
B
T
_
C
A
B
T
_
E
D
B
T
_
O
P
A
T
_
C
A
A
T
_
E
D
A
T
_
O
P
S
T
_
C
A
S
T
_
E
D
S
T
_
O
P
G
T
_
C
A
G
T
_
E
D
G
T
_
O
P
0
20
40
60
80
100
P
r
o
p
o
r
t
i
o
n

o
f

g
r
o
w
t
h

f
o
r
m
woody
succulent
herb
graminoid
climber
geophyte

Figure 11 Relative growth form composition of the microenvironments of the germinable seed bank
of the study sites. AT = Albany Thicket; BT = Buffels Thicket; GT = Gamtoos Thicket; ST = Sundays
Thicket. (CA = canopy; ED = edge; OP = open).

B
T
_
C
A
B
T
_
E
D
B
T
_
O
P
A
T
_
C
A
A
T
_
E
D
A
T
_
O
P
S
T
_
C
A
S
T
_
E
D
S
T
_
O
P
G
T
_
C
A
G
T
_
E
D
G
T
_
O
P
0
20
40
60
80
100
P
r
o
p
o
r
t
i
o
n

o
f

d
i
s
p
e
r
s
a
l

m
e
c
h
a
n
i
s
m
zoo
wind
auto

Figure 12 Relative dispersal mechanism composition of the germinable seed bank of the three
microenvironments of the study sites. AT = Albany Thicket; BT = Buffels Thicket; GT = Gamtoos
Thicket; ST = Sundays Thicket. (CA = canopy; ED = edge; OP = open).
47
Table 8 Mean (n=4) species richness (d), diversity (H), dominance (C) of the germinable seed bank
of study sites, and Sorensons co-efficient of community (CC). AT = Albany Thicket; BT = Buffels
Thicket; GT = Gamtoos Thicket; ST = Sundays Thicket. (Can = canopy; Ed = edge; Op = Open).

d

H'

C

CC

Can Ed Op Can Ed Op Can Ed Op
BT1
10.82 10.48 7.47 2.10 2.23 1.74 5.32 7.00 4.11 0.27
BT2
11.39 12.14 9.19 2.61 2.50 2.50 9.82 8.91 10.03 0.10
BT3
8.54 11.58 13.22 1.66 1.89 2.35 2.87 3.65 6.73 0.15
mean
10.25 11.40 9.96 2.12 2.21 2.20 6.01 6.52 6.96 0.17
AT1
9.11 8.83 5.17 2.63 2.00 0.96 11.66 4.41 1.69 0.15
AT2
3.99 6.62 3.45 1.36 1.90 1.54 3.02 4.22 3.74 0.12
mean
6.55 7.72 4.31 1.99 1.95 1.25 7.34 4.31 2.71 0.14
ST1
3.42 3.86 5.75 1.47 1.60 1.77 3.38 3.62 3.63 0.06
ST2
6.26 4.41 6.90 1.31 1.31 1.64 2.15 2.37 3.35 0.09
mean
4.84 4.14 6.32 1.39 1.45 1.71 2.77 2.99 3.49 0.07
GT1
5.70 6.07 6.90 1.58 1.41 1.19 2.99 2.38 1.94 0.11
GT2
4.56 6.62 9.77 1.16 1.19 1.73 2.19 1.87 3.69 0.16
mean
5.13 6.34 8.33 1.37 1.30 1.46 2.59 2.13 2.82 0.14

Buffels Thicket had the greatest species dominance as well, with the remaining three
Thicket types having lower similar values. Species dominance was greatest in the open,
with the exception of Buffels Thicket, where it was least in interclump areas.

There was a large difference in the species composition between the aboveground
vegetation and the germinable seed bank (Table 8). These differences were seen in
Sorensons coefficient (CC) which was less than 50%, with the germinable seed bank of
Sundays Thicket least similar to the vegetation, and Buffels Thicket (especially Umtiza)
the most similar.

2.4 Discussion
A clear understanding of the seed bank of a vegetation type is necessary for sustainable
management and restoration after large disturbances (Van der Valk & Pederson 1989,
Thompson 1992). If communities are overutilised and degraded, it can lead to the loss of
desirable species, as well as the establishment and subsequent dominance of unwanted
species (Florentine & Westbrooke 2004, Ellsworth et al. 2004). Communities with
48
persistent seed banks can be left alone after disturbance, but those without require
assistance to return the landscape to its natural functioning.

2.4.1 Aboveground vegetation and classification
Species richness is greatest in Sundays Thicket in the west, decreasing to Buffels Thicket
in the east. Gamtoos Thicket may not have been as rich in species as Sundays and Albany
Thicket, but it had the greatest species diversity. The three vegetation types that receive
lower rainfall roughly correspond to Valley Thicket, the structural type that has the
greatest species richness and endemism, compared to the more mesic Thicket and xeric
Arid Thicket (Vlok et al. 2003). The two sites that were not Valley Thicket (Gamtoos and
two samples of Thomas Baines) were near the boundary between Thicket and Valley
Thicket, and did not receive higher rainfall. As these two sites did not separate from
Valley Thicket, the boundaries between vegetation types in the STEP map may be
inaccurate in places.

The greater species richness and diversity may be a result of a greater number of species
co-existing together. The microclimate underneath the Thicket canopy allows drought
intolerant species to extend their range into the lower rainfall areas (Fabricius 1997).
Slow growing, drought tolerant species, including the rich succulent and geophyte flora
of Thicket, are outcompeted in mesic areas by the greater grass and herbaceous species
cover, with the resultant increase in fire frequency further preventing their establishment
and survival (Vlok et al. 2003). Thus the Thicket types that receive lower rainfall may
have a greater species richness and growth form diversity. The larger variation in species
richness of Albany and Sundays Thicket agrees with Vlok et al. (2003) who found
greatest variation in Valley Thicket.

However, Cowling (1984) and Everard (1987) found species diversity to be greatest in
Mesic Kaffrarian Thicket, rapidly declining in the west. This difference may be a result
of the three sites in this study located in Mesic Kaffrarian Thicket being situated in
mosaic Thicket types, and not solid Thicket. Thicket forms mosaics with Forest (Umtiza),
49
Grassland (Sittingbourne) and Savanna (Fort Jackson) in the east. These biomes are fire-
driven (Low & Rebelo 1998), which may limit the persistence of Thicket species that do
not tolerate fire (Kerley et al. 1995), causing a decrease in species diversity in these
Thicket mosaics. It may also be due to a different sampling method being used, with both
Cowling (1984) and Everard (1987) using fixed quadrats of 100 m
2
, and describing
species richness based on area sampled, compared to this study that estimated the cover
of the species in the immediate surrounding area.

The failure of the sites to conform phytosociologically to Everard (1987) resulted in the
use of the STEP classification scheme. The three vegetation types studied here showed
less separation of phytosociology compared to Everards (1987) DECORANA analysis of
the vegetation. Mesic Kaffrarian Thicket remained separate from the remaining study
sites, and like Everard (1987) had the greatest compositional variation, but no clear
separation existed between Xeric Kaffrarian Thicket and Mesic Succulent Thicket,
evident in Everard (1987), especially in the Gamtoos River valley and Port Elizabeth in
the west. All of Everards (1987) XKT sites were situated along the coast, unlike the
sampling sites in this study that were situated more inland, and should be considered
Dune Thicket (Low & Rebelo 1998, Vlok et al. 2003). The remainder of his Xeric
Kaffrarian Thicket vegetation type would be considered Valley Thicket in the river
valleys and Thicket between the valleys (sensu Vlok et al. 2003). Inland Xeric Kaffrarian
Thicket could be as succulent as Mesic Succulent Thicket and therefore more similar to it
than Dune Thicket.

The hierarchical classification of Thicket in STEP (Vlok et al. 2003) allowed the
sampling sites to be classified into biogeographical regions that better explained the
relationships between sites. Sundays Thicket and Gamtoos Thicket overlap in ordination
space, as they are geographically close to each other. The structure and diversity of
Gamtoos Thicket is intermediate between Sundays and Groot Thicket, resulting in diffuse
patterns in species gradients, and neighbouring vegetation types enriching each other
(Vlok & Euston-Brown 2002). Sundays Thicket west of the Swartkops River contains
50
more species similar to Gamtoos Thicket, causing this area to be a transition zone (Vlok
& Euston-Brown 2002).

However, the study sites did not separate in ordination space according to STEPs
structural units. The Thicket units that were sampled outside of Buffels Thicket (Gamtoos
and Albany Thicket) were very near the boundary between Valley Thicket and Thicket,
and many similarities could therefore be expected. Rainfall was very near the lower
boundary for the rainfall range for Thicket (Vlok & Euston-Brown 2002), further
suggesting the presence of more xeric species. The character species of Thicket
(Euphorbia triangularis, Olea europaea subsp. europaea, Rhus pallens, Scutia myrtina)
occurred at Gamtoos as well as Loerie, with the exception of E. triangularis. The two
samples classified as Thicket at Thomas Baines (AT1) contained only Scutia myrtina,
whereas all four species were present in the samples classified as Valley Thicket.

2.4.2 Seedlings and the microenvironment
Seedlings were found at all sites, although in very low numbers. Both the number of
species and number of seedlings decreased with an increase in xeric conditions. Seedlings
of woody species dominated in the mesic east, whereas succulent seedlings are more
important in the xeric west. The regeneration of species by seedlings is much greater in
the mesic sites of Buffels Thicket compared to the more xeric Sundays Thicket. This
trend is also identified within Albany and Gamtoos Thicket, where the more mesic sites
(Thomas Baines and Gamtoos) had greater seedling richness than the other sites. It can
therefore be assumed that an increase in annual rainfall will result in an increase in the
diversity of species that successfully regenerate through sexual reproduction (Vlok &
Euston-Brown 2002, Sigwela 2004). The low number of seedlings may also be due to
sampling taking place in autumn and winter. In the west, bimodal rainfall with peaks in
spring and autumn will result in most seeds expected to germinate in these seasons.
However, in the east, there is a greater proportion of summer rainfall, with winter being a
drought season (Vlok et al. 2003). However, the assumption that Thicket regeneration is
occurring when seedlings are found must be made tentatively. Although seedlings are an
51
indicator of successful sexual reproduction of species (Fenner & Thompson 2005),
seedling survival may be low (De Ridder 2006), resulting in minimal recruitment into the
canopy.

Seedlings of only a few species in the canopy at the various sampling sites were found.
Succulents, especially arborescent species, dominate the seedling composition of
Sundays Thicket. This is similar to Midgley & Cowling (1993). Succulents are able to
establish in dry conditions that may cause mortality in other growth forms as they are
capable of Crassulacean Acid Metabolism (CAM), allowing them to survive and grow in
low water conditions (Osmond et al. 1981).

Seedling patterns identified in this study is similar to previous findings (La Cock 1992,
Sigwela 2004), as seedlings were observed in all vegetation types in the study. The total
number of species found is greater than that of Sigwela (2004) (n=11) and Midgley &
Cowling (1993) (n=25). This may be due to more mesic conditions being experienced
compared to Sundays Spekboom Thicket (Sigwela 2004), where mean annual rainfall is
below 400 mm.

The lower light and temperatures experienced underneath the canopy of Thicket is a
widely found phenomenon (Sigwela 2004), providing a microclimate beneficial for the
survival of seedlings, especially in the more xeric areas. It would be expected that most
seedlings will occur at the edge of Thicket, where light levels are intermediary between
the canopy and in interclump areas (De Ridder 2006). This, accompanied by the
ameliorated temperatures and greater amounts of litter, will promote the emergence and
establishment of seedlings.

The litter layer in a community may either stimulate or inhibit the germination of seeds
(Vasquez-Yanes & Orozco-Segovia 1993). Litter can act as a nursery by holding enough
moisture for the germination of large seeds. It can also act as a barrier to seedling
emergence for species that do not have the resources to penetrate through the layer
(Ellsworth et al. 2004), as well as changing the ratio of red: far red light at the soil
52
surface, preventing germination of seeds that are light-sensitive (Vasquez-Yanes et al.
1990). In Thicket, litter is beneficial as a source of nutrients (La Cock 1992) and will not
prevent seedling emergence, as most species are large-seeded (Watson 2002), and should
be able to penetrate the litter layer (Vasquez-Yanes et al. 1990).

2.4.3 Characteristics of seed bank
The seed bank of all Thicket vegetation types in this study were dominated by grasses
and herbs. Woody species made a larger contribution to the relative proportion of growth
forms in Buffels Thicket, whereas succulents were more important in Sundays and
Albany Thicket. In Gamtoos Thicket, woody species were sub-dominant underneath the
canopy, as opposed to succulents which were sub-dominant in the open. Short-lived
herbs that are required to replace their populations regularly to survive form persistent
seed banks (Murdoch & Ellis 1992). Many herbs and some grasses found in this study are
weedy or pioneer species, and are likely to be able to form persistent seed banks,
dominating the seed bank of Thicket. The dominance of grasses and herbs is similar to
previous studies in Thicket (Pierce & Cowling 1991, Sigwela 2004) as well as similar
forest systems (Bossuyt et al. 2005).

Seed densities in the seed bank were low, similar to other stable vegetation types
(Garwood 1989). The increased amount of seeds in the germinable seed bank of Buffels
Thicket is a reflection of greater species richness, as no single species dominated the seed
bank. Pierce and Cowling (1991) found that the seed bank in Thicket was dominated by a
single herb, Lobelia erinus. This is comparable to forest communities, where seed banks
are usually dominated by a single species, often a pioneer tree (Garwood 1989). The
density of seed banks may be highly underestimated by the seedling emergence
technique, as specific germination cues may not be met (Simpson et al. 1989). However,
as this was a community level study with a high number of species, determining the size
and composition of the seed bank by seedling emergence is preferable as direct counting
is tedious and requires viability testing (Simpson et al. 1989). The use of both methods
53
would not have been feasible, due to the large number of samples and high species
diversity (Pierce & Cowling 1991).

In Thicket, the composition of the seed bank seems to be a result of a number of factors,
including seed dispersal, seed predation and dormancy. In communities where most
species have large fruits, such as Thicket, a small seed bank would be expected, as
granivores target larger seeds, thus having a large influence on seed banks (Garwood
1989). Rodents remove more seeds in semi-arid systems than ants (Anderson &
MacMahon 2001, Sigwela 2004). Granivory is greater in untransformed than transformed
Thicket (Sigwela 2004), and in dense bushclumps compared to open grassy areas
(Watson 2002). Therefore granivores will result in a larger decrease in the number of
large canopy species seeds underneath the canopy, as opposed to clearings or openings.
As large-seeded species are usually vertebrate-dispersed (Garwood 1989), few seeds
would end up in clearings, resulting in the complete loss of species with large seeds from
the seed bank.

No clear patterns between the seed banks of interclump areas, compared to the edge and
underneath the canopy in terms of species richness and abundance of seeds in the seed
bank. In the coastal shrub thickets of North America, seed banks of clearings are richer
than underneath the canopy, and contain more woody species (Crawford & Young 1998).
Intermediate seral stages with intermediate levels of cover contain seed bank with highest
species richness (Crawford & Young 1998). The lack of difference in the germinable
seed bank of openings and underneath the canopy may be explained by the dominance of
grasses and herbs that have small seeds, as small-seeded species are more frequently
observed and more broadly distributed over space than large-seeded species (Guo et al.
1998).

Anemochory was the dominant dispersal mechanism throughout the vegetation cover
gradient. Zoochory was relatively more important underneath the canopy than in
openings, except in Sundays Thicket where the greatest contribution was to the edge of
Thicket clumps. The dominance of anemochorous species is caused by the seed bank
54
being dominated by wind-dispersed weed species, a beneficial adaptation as wind-
dispersed species are generally dispersed further than the other mechanisms (Fenner &
Thompson 2005). As both birds and mammals prefer the safety of the dense cover
(Kollman 2000, Watson 2002), their relative dominance in the under canopy environment
of Thicket is not unexpected. In arid areas, the distribution of seeds in the seed banks is a
result of different dispersal mechanisms. In Deserts, seed banks have greater species
richness, as well as being denser underneath shrubs compared to open areas (Guo et al.
1998). This was due to the shrubs catching and collecting wind- and self-dispersed
species. In Grassridge Bontveld, a mosaic Thicket vegetation type, zoochory dominated
the bushclumps, compared to the open areas dominated by grasses or succulents, where
anemochory and autochory were the dominant dispersal mechanism, respectively
(Watson 2002).

It could be incorrect to assume that all wind dispersed species found underneath the
canopy were dispersed into that environment by the wind. Pakeman et al. (2002) found
that species that have adaptations to other mechanisms of dispersal, such as wind,
survived ingestion and passage through the gut of an animal, germinating in large
numbers from animal dung. Small round seeds were the dominant form of seeds that
germinated from the dung, whereas large or elongated seeds did not germinate. Hard
seeds germinated as well.

2.4.4 Similarity between seed bank and aboveground vegetation
There was a low similarity between the germinable seed bank of and the aboveground
vegetation of Thicket. This was due to the dominance of herbs and grasses, and the
absence of long climax woody tree and succulent species. Dominant Thicket species do
not form persistent seed banks, and mostly regenerate through the production of ramets
(Midgley & Cowling 1991). As a subtropical, evergreen vegetation type that does not
experience large periodic disturbances, but is surrounded by a relatively disturbed
landscape, the seed bank of Thicket will fall somewhere between that of temperate and
tropical Forests. It is similar to temperate Forests, as few climax species accumulate
55
persistent seed banks (Pickett & McDonnell 1989, Garwood 1989, Thompson 1992). The
seed bank is dominated by pioneers such as grasses and herbs, which are usually species
that are not found in the existing vegetation community but grow somewhere else, and
were dispersed into the forest (Baskin & Baskin 1989) like temperate Forests, but tend to
be quite small, similar to tropical dry Forests. Most persistent seeds are small, with the
exception of hard seeds and legumes (Baker 1989).

It is common for the seed bank of a community to reflect a pioneer successional
community (Crawford & Young 1998). In the woodlands and bushlands of Ethiopia, the
seed bank was dominated by herbs and grasses while many woody species in the
vegetation did not occur in the seed bank (Mengistu 2005). The dominance of grasses and
herbs is consistent with previous studies into the seed bank of Thicket (Pierce & Cowling
1991, Sigwela 2004).

2.4.5 Implications for regeneration through seed bank and seedlings
The contribution that a seed bank makes to regeneration is mainly a product of the size
and shape of gaps in the vegetation. In regularly disturbed sites, as well as sites that
experience large-scale disturbances, the seed bank will make a large contribution to the
composition of the vegetation, while in small gaps and stable communities, most
regeneration will be vegetative (Luzuriaga et al. 2005). Historically, megaherbivores
created paths through Thicket (Kerley et al. 1995), but any gaps would be quickly filled
with vegetative sprouts. In addition, gaps in Thicket are not preferential sites for the
establishment of seedlings, due to abiotic stresses (Sigwela 2004). Therefore the seed
bank will more than likely not make a considerable contribution to the regeneration of
Thicket, especially as it lacks most of the dominant woody species found in the
aboveground vegetation.

The limited number of canopy species in the seed bank, were concentrated in the more
mesic Thicket. Acacia karroo, Buddleja saligna, Cussonia spicata, Diospyros
dichrophylla, Grewia occidentalis, Rhus sp., Lycium ferocissimum, Phyllanthus
56
verrucosus, Putterlickia pyracantha and Sideroxylon inerme were recorded in the seed
bank. Many of these species, with the exception of S. inerme and C. spicata, could be
considered to be pioneer species, and are usually some of the first woody species to
colonise disturbed areas (Hall 2000). Even though Sigwela (2004) suggests that Thicket
species do not have recalcitrant seeds and thus cannot form persistent seed banks, it was
observed in this study that some species only emerged after a couple of months, or even
as long as a year. As traits that are selected by zoochory are similar to those found in
species with persistent seed banks (Pakeman 2002), it is possible that some species do
have seeds capable of dormancy that persist in the soil for greater than a year. Other
factors, such as the lack of safe sites or granivory, could be the cause of the lack of
regeneration through seedlings.

Areas where seeds collect are usually safe sites, where conditions are favourable for
seedling establishment (Kinloch & Friedel 2005b). Overgrazing and transformation of the
landscape decreases the number of safe sites for seedling establishment (Lechmere-
Oertel 2003, Sigwela 2004) and places where seed can accumulate (Kinloch & Friedel
2005b). Dead and fallen woody species act as nursery sites for seedling regeneration, and
fuelwood harvesting may further limit safe sites (Sigwela 2004). The removal of nurse
species and other trees will result in the further loss of perches for birds, as well as habitat
for mammals, preventing the dispersal of zoochorous species (Watson 2002, Lechmere-
Oertel 2003).

Sigwela (2004) suggests that seedling regeneration in Thicket tends to be linked to
climatic variation. Xeric conditions favour vegetative regeneration, whereas mesic sites
have greater numbers of seedlings, therefore causing sexual reproduction to become
dominant. The survival and establishment of seedlings is probably episodic and
infrequent, only occurring during very wet seasons (Stuart-Hill & Danckwerts 1988,
Sigwela 2004).

Seedling emergence and survival is highest at the edge of Thicket (De Ridder 2006) and
other forest (Lpez-Barrera et al. 2006) and shrubland ecosystems (Arrieta & Surez
57
2006), as well as under shrubs in more open areas (Cowling et al. 1997, Arrieta & Surez
2006) where intermediate light and moisture conditions, as well as competition occurs.
Even though seedling emergence can be greater underneath the canopy, seedling survival
was highest at the edge of an European woodland (Arrieta & Surez 2006). This suggests
that seedling abundance is more likely a product of seedling survival than successful
germination at the edge of Thicket clumps. The preferential environment at the edge of
Thicket clumps will lead to their expansion, until they finally merge. The loss of
megaherbivores from the landscape will cause Thicket to become denser, leading to a
decrease in the edge environment, and a predicted decrease in seedlings.

The presence of the non-Thicket tree species Celtis africana and Vepris lanceolata
indicates that there is dispersal occurring into Thicket from neighbouring Forest patches.
However, as these plants are absent from the canopy, it is assumed that rainfall is not
high enough for the successful establishment of these species. If climate change results in
an increase in rainfall, there may be a biome shift from Thicket to Forest in the mesic
areas of the east.

Even though there is very little similarity between the seed bank and aboveground
vegetation, with few canopy species present, the seed bank could still play an ecological
role in the stabilisation of the soil after a disturbance, preventing soil erosion (Sigwela
2004). Large pioneer shrubs that occur outside of Thicket clumps but do form part of the
seed bank may act as a perch for birds (Cowling et al. 1997, Watson 2002), and provide
cover for bird-dispersed species. Without this initial cover, the canopy will not become
established.

2.4.6 Conclusion
The seed banks of mesic Thicket types are dominated by herbs and grasses, allowing the
first hypothesis to be accepted. The regeneration of Thicket is not dependent on its
germinable seed bank, and any program with the intention of restoring Thicket will not
be able to rely on it either. Woody species are only more relatively important underneath
58
the canopy of Buffels and Gamtoos Thicket as opposed to openings, rejecting the second
hypothesis. Therefore the lack of seedlings in the open observed by other authors (La
Cock 1992, Sigwela 2004) cannot be a result of a lack of dispersal, but must therefore be
a result of poor seedling survival. If dispersers are present in the landscape and have not
been removed by human activities, dispersal should occur into new open areas.

Although anemochory was the dominant dispersal mechanism underneath the Thicket
canopy, as well as in the interclump areas, zoochory was relatively more important to the
germinable seed bank below the canopy. The third hypothesis that zoochory is the
dominant dispersal mechanism underneath the canopy is therefore only partially rejected.
The relative importance of zoochory indicates that animal-dispersed species are
preferentially dispersed to Thicket clumps. As woody species are predominantly
zoochorous (Watson 2002), and are not limited in the germinable seed bank to Thicket
patches, higher granivory underneath the canopy may be limiting seedling establishment
in Thicket. It can be provisionally assumed that a lack of seedling survival may limit
regeneration through seedlings in the open, but granivory limits it underneath the canopy
of Thicket.

Finally, the last hypothesis that seedlings are more abundant in Thicket types that receive
higher rainfall is accepted, as Buffels Thicket had both the greatest seedling abundance
and richness. Seedlings were found throughout the study area, but in low numbers,
limiting their possible contribution to the regeneration of Thicket.
59
Chapter 3 Dormancy and germination in Thicket species
3.1 Introduction
An understanding of the germination requirements of species is critical for the
understanding of natural systems, ranging from the regeneration niche of individual
species (Vandvik & Vange 2003), to the assemblage rules and succession of communities
(Parker et al. 1989), and the restoration and rehabilitation of ecosystems (Bell et al.
1993). Ecologists that work on the germination requirements of a species are challenged
to define the environmental conditions necessary to bring about changes in seeds that
result in a release from dormancy and to correlate these with factors that promote
dormancy break and emergence from the soil in nature (Teketay & Granstrm 1997,
Baskin & Baskin 1998, Burke et al. 2003).

Germination begins when the dry seed takes up water, a process known as imbibition
(Bewley & Black 1994). This leads to a rapid rise in respiration, the mobilisation of
nutrient reserves and the initiation of growth in the embryo (Fenner & Thompson 2005).
The process is completed when the testa (seed coat) ruptures and the embryo emerges as
the plumule or radicle, allowing photosynthesis to begin (pik & Rolfe 2005). Once the
seed begins to germinate, it is committed to growth or death as germination is irreversible
(Fenner & Thompson 2005).

However, some seeds do not germinate immediately when exposed to water (Rolston
1978). This may be due to quiescence or dormancy. A seed is said to be quiescent if the
minimum requirements for germination are absent, preventing the growth of non-dormant
seeds or organs (Murdoch & Ellis 1992, Foley 2001). A seed in a brown paper bag at a
laboratory desk would be quiescent (Baskin & Baskin 1998). For germination to occur,
the chemical environment must be right (Bewley & Black 1994). Water must be readily
available, there must be a good supply of oxygen for respiration, and any chemicals that
prevent germination must be absent (Edwards et al. 1988). The physical environment is
also important with the correct temperature and quality and quantity of light being
present (Bewley & Black 1994).
60

3.1.1 Seed Dormancy
Dormancy differs from quiescence in that it is a characteristic of the seed and not the
environment and defines what conditions should be met to make the seed germinate
(Vleeshouwers et al. 1995). A seed is said to be dormant when it fails to germinate even
though the environmental conditions required for germination (water, light etc.) are
present (Murdoch & Ellis 1992; pik & Rolfe 2005). Dormancy can be defined as a
block within the seed that prevents germination (Vleeshouwers et al. 1995) and can result
from the absence of signals that are required for growth or the presence of inhibitory
signals that prevent growth, with the combination of the two also being possible (pik &
Rolfe 2005). These signals may either originate from the seed coat (seed-coat imposed
dormancy) where the seed coat acts as a physical or chemical barrier (Rolston 1978) or
from the embryo (embryo-imposed dormancy) where plant hormones play a role (Ernst et
al. 1988).

It is commonly thought that the function of dormancy in seeds is to prevent germination
during periods that are not conducive to germination and establishment. However, seeds
will not germinate during the dry season due to the lack of imbibition, or during the cold
season as a result of relatively high temperatures being required (Vleeshouwers et al.
1995). Dormancy is only necessary when the conditions for germination are met but the
probability of seedling survival and growth is negligible (Vleeshouwers et al. 1995), and
is only beneficial if it improves plant fitness by allowing germination to take place in
circumstances that would result in the highest probability of subsequent survival and
reproduction (Vandvik & Vange 2003). Therefore specific cues are required, not random
and unpredictable ones (Fenner & Thompson 2005).

Dormancy increases the chances of the successful establishment of seeds, by spreading
germination over space and time (Rolston 1978, Foley et al. 2001). Germination is
spread over time, by some species producing seeds that have different degrees of
dormancy, which is often manifested in their appearance (Campbell & Matthewson
61
1992). This phenomenon is known as polymorphism and allows seedlings to emerge at
irregular intervals resulting in reduced competition and a greater chance of survival
(Forsyth & Brown 1982, Dubey & Pandey 1988). The breaking of dormancy may be
reliant on an environmental factor which itself has a time distribution, such as light and
temperature. Seeds that require a period of high temperatures to break dormancy ensure
that germination will take place during the cooler time of the year, increasing seedling
survival (Bell et al. 1993, Vleeshouwers et al. 1995). Germination may also occur over
space, as many species have dormancy which is terminated by light, with red light being
the most effective. This allows seeds to germinate in the open where competition is less
and not under a canopy where light is poor in the red component, or not at depths in the
soil that will prevent seedling emergence (Teketay & Granstrm 1997, Pearson et al.
2002, Vandvik & Vange 2003).

3.1.2 Types of Seed Dormancy
There are different types of dormancy that have been classified by various authors
(Harper 1957, Nicolaeva 1977, Baskin & Baskin 1998). Harper (1957) introduced the
terms innate, induced and enforced dormancy when describing dormancy that develops
on the mother plant, dormancy that is induced after shedding and dormancy that is
enforced by unsuitable environmental conditions, respectively (Murdoch & Ellis 1992).
Innate dormancy occurs when a trait of the seed itself blocks germination (Burke et al.
2003). It may be as simple as the embryo still being immature or even undifferentiated
when it is shed from the mother plant, requiring further development before germination
occurs (Harper 1957). Germination may also be blocked by some physiological aspect of
the seed, such as a hard-seed state, irrespective of embryo maturity (Murdoch & Ellis
1992). Induced dormancy occurs when dormancy can be brought about in both dormant
and non-dormant seeds once they have been shed from the parent plant. This occurs when
moist or fully imbibed seeds are kept in environmental conditions that do not promote
germination, or are insufficient to break dormancy in the seed (Teketay & Granstrm
1997). Enforced dormancy occurs when there is an environmental restraint to
62
germination (Harper 1957). However, this type of dormancy is now more commonly
accepted as quiescence (Fenner & Thompson 2005).

As dormancy is now considered a trait of the seed and not the environment, this
classification is no longer in favour as both induced and enforced dormancy is caused by
incorrect environmental conditions (Fenner & Thompson 2005). The terms primary and
secondary dormancy are now preferred (Foley 2001). Primary dormancy occurs when
freshly matured seeds fail to germinate when incubated over a range of conditions (Foley
2001). This is equivalent to innate dormancy. Non-dormant seeds may enter dormancy if
environmental conditions become unfavourable for germination (Vleeshouwers et al.
1995). This is known as secondary dormancy, and is similar to induced dormancy, except
for the fact that there are no reported cases of seeds that are non-dormant at maturity
being induced into dormancy (Baskin & Baskin 1998).

According to Fenner and Thompson (2005), there are essentially three types of
dormancy. These are physical, morphological and physiological dormancy. Seeds with
physical dormancy, commonly known as hard seeds, have testas or pericarps that are
impermeable to water, preventing the embryo from imbibing water and germinating
(Rolston 1978, Ernst et al. 1988). They generally have hard seed coats that contain a
palisade layer of lignified cells, which are often impermeable (Li et al. 1999). Seeds that
lack a well-differentiated seed wall are protected by the fruit wall that may be
impermeable to water, imposing physical dormancy, such as Rhus lancea in the family
Anacardiaceae (von Teichman 1989). Seed impermeability may also be a result of the
natural seed openings, which include micropyle, lens, hilum and chalazal areas,
becoming impermeable to water (Rolston 1978, Li et al. 1999, Baskin 2003).

Morphological dormancy occurs when the embryo of a seed is still immature at the time
of shedding from the parent plant, and a period of growth and/or differentiation is
required before germination will take place (Fenner & Thompson 2005). This may be a
result of either the embryo being undifferentiated, or underdeveloped. These seeds tend to
be very small, and are classified as micro or dwarf seeds (Baskin & Baskin 1998).
63
However, Foley (2001) does not recognise this as a form of dormancy, as growth is
required for germination, and not the breaking of dormancy.

Physiological dormancy is caused by a physiological inhibiting mechanism of the seed
that prevents radicle emergence (Baskin & Baskin 1998). Structures that cover the
embryo (endosperm seed coats, fruit walls and husks) may also play a role in preventing
germination (Pearson et al. 2003), by mechanically or chemically preventing the
diffusion of oxygen into the seed (Kahn 1977, Forsyth & Brown 1982), by preventing
chemical inhibitors from leaching out of the seed, and by mechanically constraining the
growth of the embryo (Mayer 1977, Hemmat et al. 1985). There are various causes of
physiological dormancy, which may be environmental, such as a specific light (Forsyth &
Brown 1982, Teketay & Granstrm 1997), temperature (Hemmat et al. 1985, Teketay &
Granstrm 1997, Burke et al. 2003, Vandvik & Vange 2003), or chemical requirements,
including smoke (Brown 1993, Brown & Van Staden 1997, Lloyd et al. 2000) and
nutrients (Bell et al. 1993). Chemicals inhibitors may be found in the embryo, or the
endosperm and seed coat (Forsyth & Brown 1982, Teketay & Granstrm 1997), as well
as structures that are dispersed with some seeds (Campbell & Matthewson 1992). These
inhibitors either have to be deactivated or leached out to allow germination (Campbell &
Matthewson 1992). Unlike the two other types of dormancy, physiological dormancy is
reversible with the seed able to enter into secondary dormancy, allowing a more flexible
response to the environment (Baskin & Baskin 1998).

Combinations of these three dormancy types also exist, with morphophysiological
(morphological and physiological) dormancy being the most common (Baskin & Baskin
1998). Physical and physiological dormancy is rare while physical and morphological
dormancy is impossible (Fenner & Thompson 2005).

A simpler classification, based on the mechanisms or location of constraints to
germination, is separating coat-imposed from embryo dormancy, according to whether
dormancy is imposed by the seed coat, or factors within the embryo (Foley 2001). As this
64
is an ecological study and it was not possible to determine whether the seed coat imposes
physical or physiological dormancy, this classification will be used in this study.

The different dormancy types control the habitats and climates that species possessing
them can occupy, as well as their preferred regeneration niche. Seeds with morphological
or morphophysiological dormancy need to be imbibed for embryo differentiation and/or
growth to occur and therefore generally occur in regions with moist seasonal climates,
including woodlands and damp Grasslands (Fenner & Thompson 2005). Seeds with
physical dormancy do not need to be imbibed when dormancy is broken and are common
in habitats with distinct dry seasons such as tropical deciduous Forests, Savannas, hot
Deserts, steppes and Fynbos where high temperatures, large temperature fluctuations and
fire occur (Rolston 1978, Bell et al. 1993). Many forest species, most of them pioneers,
have physical or physiological dormancy that is broken by large fluctuations in
temperatures or high irradiance, to ensure successful establishment in forest gaps
(Teketay & Granstrm 1997, Pearson et al. 2002).

3.1.3 Factors affecting the germination of Thicket species
There is a large contrast between the soil environment underneath Thicket clumps and
open areas, especially in xeric areas. The soil underneath the canopy is nutrient-rich
(Mills & Fey 2002, Sigwela et al. 2003), has a deep litter layer that retains moisture and
dampens temperature changes (Lechmere-Oertel 2003) and contains mycorrhizal fungi
(Allsopp et al. 1996). Soil moisture is relatively higher (Mills & Fey 2004) and pH is
lower compared to open areas in the Thicket (Lechmere-Oertel 2003). Open areas are
associated with a loss of topsoil and cover, particularly litter, in xeric areas (Lechmere-
Oertel 2003) and is highly compact, physically preventing the establishment of seedlings
(Todkill 2001). Greater evaporation occurs in the open, drying out the soil and causing
crusting (Mills & Fey 2004). The lack of cover reduces mychorrhizal abundance, except
where a high grass cover is maintained in openings (Allsopp et al. 1996). In mesic areas,
the greater grass and herb cover in the open will reduce temperature and improve soil
moisture retention, improving conditions for germination.
65

There is a reduction in light intensity below the Thicket canopy (Chapter 3, Sigwela
2004) accompanied by a decrease in the red:far-red light ratio. Temperatures are also
significantly lower underneath the canopy (Lechmere-Oertel 2003) and temperature
fluctuations are likely to be less than in the open. Minimum temperature reaches below
5C for all Thicket types, with the exception of Dune Thicket (8 C), and maxima may
reach above 50C in xeric inland areas (Vlok et al. 2003).

Many Thicket species have fleshy fruits and are endozoochorous (Cowling1983, Bruton
1999, Watson 2002). The passage through the gut of an animal removes the flesh from
the seed, together with any chemical inhibitors that occurred within it (Sigwela 2004).
The seed coat may also be affected and made permeable to water, allowing the seed to
imbibe water (Rolston 1978). Species adapted to endozoochory generally have small,
round seeds, and have physical dormancy.

Most Thicket canopy species have a tropical origin (Cowling 1983). Many tropical forest
species have fleshy, non-dormant fruits that germinate immediately, and in shade (Van
der Pijl 1972, 1984, Garwood 1989). They are generally only viable for a few months,
and lack any dormancy mechanisms (Garwood 1989). However, dormancy is common in
the dry Afromontane forests of Ethiopia (Teketay & Granstrm 1997), where 60% of
species had some degree of dormancy, and only one species was found to be recalcitrant.
Dominant dormancy mechanisms are physical in species adapted to fire, and
physiological if broken by light or temperature changes (Teketay & Granstrm 1997).

The dormancy mechanisms of Thicket species, as well as the related dormancy breaking
mechanism should be known in order to fully understand regeneration of Thicket, as well
as individual regeneration niches of Thicket species. If seed germination is sensitive to
the environment, seedling survival can be greatly increased, as certain periods of time
and locations that could result in greater mortality can be avoided (Vandvik & Vange
2003). The aim of this chapter is to further the understanding of the germination
requirements of Thicket species, and relate these requirements to their regeneration niche.
66

The objectives of this chapter are to:
1. Identify the dominant dormancy mechanisms of woody and succulent species in
Thicket;
2. Predict the regeneration niches of species, based on their dormancy mechanisms.

Hypotheses:
H1: Seeds with thick and/or hard seed coats will have seed coat-imposed dormancy, and
mechanical scarification and/or soaking in hot water will promote their germination;
H2: Seeds that have thin seed coats will have embryo dormancy, and germination may be
promoted by either cold stratification or shade, or prevented by nutrient-poor sand.

The species selected for this study was based on Everards (1987) characteristic species
for Thicket vegetation types, as well as the availability of large quantities of seeds from
the seed suppliers. Unfortunately, the age of the seeds could not be supplied by the
supplier. Seeds were received in February 2008 with their flesh removed, and pre-treated
with a fungicide. They were stored in paper bags in a cool dry environment until needed.
Not all species were exposed to all the treatments as the number of seeds available was
insufficient for some species. A short description of the study species is given in Table 9.
Information was gathered from Palmer & Pitman (1972), Coates-Palgrave (2002) and
Germishuizen and Meyer (2003). The collection location, treatments and seed
characteristics are presented in Table 10.

Seeds were scarified with sand paper for the mechanical scarification treatment. Nicking
was more appropriate for Olea europaea and Mystroxylon aethiopicum, as the flesh had
been removed by the seed supplier and sand paper was found to be ineffective against
their thick seed coats. To determine the effect of hot water, seeds were left in hot tap
water for 24 hours. These three treatments were done to identify whether seed coat
prevents germination.
67
Table 9 Ecological characteristics and distribution of study species.
Species Family Growth from Vegetation type Distribution Dispersal
Acacia karroo Hayne
Fabaceae:
Mimosoideae
Small tree, up to 15
m tall Woodland, arid watercourses
Namibia to SW Cape, up to
Zimbabwe mammal
Aloe africana Mill. Asphodelaceae
Small succulent tree,
up to 4m tall Dense woodland, Thicket Western EC wind
Brachylaena elliptica (Thunb.) DC. Asteraceae
Shrub or small tree,
4-10 m tall
Forest margins, coastal and riverine
woodland, grassland EC and KZN coastal areas wind
Burchellia bubalina (L.f.) Sims Rubiaceae
up to 10 m tall Evergreen forest, grassland
Limpopo to SW Cape along
coastal areas bird
Carissa bispinosa (L.) Desf. ex
Brenan Apocynaceae
up to 5 m tall
Evergreen forest, dry woodland, dune
scrub
Namibia; SW Cape to
Mozambique and Zimbabwe
along eastern seaboard
bird,
mammal
Cussonia paniculata Eckl. & Zeyh.
var. paniculata Araliaceae
Short tree, up to 5 m
tall
Dry woodland, dense woodland, mountain
slopes
Limpopo and Botswana to SW
Cape bird
Cussonia spicata Thunb. Araliaceae
Short tree, 3-10 m
tall
Open woodland, forest margins, mountain
slopes
Zimbabwe and Mozambique to
SW Cape along coastal areas bird
Diospyros dichrophylla (Gand.) De
Winter Ebenaceae Shrub, 2-3 m tall
Coastal scrub, grassland, open woodland,
forest margins
SW Cape to Swaziland along
coastal areas; Limpopo mammal
Euclea natalensis A.DC. subsp.
natalensis Ebenaceae
up to 12 m tall
Coastal scrub, dry woodland, evergreen
forest, forest fringes, swamp
Widespread; Ethiopia to SW
Cape bird
Euclea racemosa Murray Ebenaceae
up to 12 m tall Dune scrub, coastal forest Namibia to EC; KZN bird
Euphorbia mammillaris L. Euphorbiaceae
Succulent Shrub, up
to 0.5 m tall Dense woodland, karroid scrub WC and EC self
Euphorbia tetragona Haw. Euphorbiaceae
Succulent tree, up to
7 m tall
Dry hillsides and valley slopes, thorn
scrub, scrub forest coastal areas of EC and KZN bird, self
Grewia occidentalis L. Tilicaeae
up to 6 m tall
Evergreen forest, open woodland, thicket,
coastal scrub
SW Cape to Zimbabwe and
Mozambique bird
Grewia robusta Burch. Tilicaeae
up to 3 m tall Karroid scrub, thicket
Western EC and inland WC and
NC bird
Gymnosporia capitata (E.Mey. ex
Sond.) Loes. Celastraceae Shrub, 2-3 m tall Dense woodland, river valleys WC and EC bird
Hippobromus pauciflorus Eckl. &
Zehy. Sapindaceae Small tree, 3-5 m tall Riverine fringes, forest margins
Limpopo to SE Cape along
coastal areas bird
Maytenus heterophylla Celastraceae
Shrub to small tree,
up to 6 m tall Evergreen forest, woodland, karroid scrub
Cape bird
Mystroxylon aethiopicum (Thunb.)
Loes. subsp. aethiopicum Celastraceae
up to 12 m tall
Evergreen forest, riverine fringes, open
woodland
Cape bird
Olea europaea L. subsp. africana
(Mill.) P.S.Green Oleaeceae
up to 18 m tall Riverine fringes, open woodland
Widespread; Zambezi River to
SW Cape
bird
cont.
68
Species Family Growth from Vegetation type Distribution Dispersal
Pappea capensis Eckl. & Zeyh. Sapindaceae Small tree, 3-9 m tall Open woodland, riverine fringes
Zimbabwe to SE Cape; Karoo to
S Namibia bird
Ptaeroxylon obliquum (Thunb.)
Radlk. Rutaceae
Shrub to large tree,
up to 20 m tall Evergreen forest, open woodland, thicket
Widespread; Tanzania to SE
Cape; Namibia and Angola wind
Putterlickia pyracantha (L.) Szyszyl. Celastraceae
up to 6 m tall
Dense woodland, thicket, fynbos, dune
forest
Coastal areas of SW Cape to
KZN bird
Schotia brachypetala Sond.
Fabaceae:
Caesalpinoideae Tree, up to 16 m tall
Open woodland, scrub forest, riverine
fringes EC coast up to Zimbabwe mammal
Scutia myrtina (Burm.f.) Kurz Rhamnaceae
up to 8 m tall
Coastal forest, forest margins, dense
woodland
Widespread; Uganda to SW
Cape bird
Zanthoxoylum capense (Thunb.)
Harv. Rutaceae
Small tree, up to 10
m tall Dry woodland, mountain slopes
SE Cape to Mozambique and
Zimbabwe bird
69
Table 10 Seed source, mean size (n = 10), mean weight (n = 10*) and relevant treatments of study
species. (MP = Mpumalanga, SC = southern Cape, EC = Eastern Cape, ex hort = horticultural stock).
(treatments: MS = mechanical scarification, HW = hot water, CS = cold stratification, Sa = sand, Sh
= shade).
Species seed source
seed
length
seed
width
seed
weight
Treatments

(mm) (mm) (mg) MS HW CS Sa Sh
Acacia karroo MP 6.04 1.66 41.54 1 1 1 1 1
Aloe africana SC 3.84 0.88 2.51 1 1 1 1 1
Brachylaena elliptica EC 3.48 0.59 0.66 1 1 1 1 1
Burchellia bubalina EC 2.98 0.71 2.32 1 1 1 1 1
Carissa bispinosa EC 7.66 2.03 23.37 1 1
Cussonia paniculata ex hort 4.83 2.65 18.71 1 1 1 1 1
Cussonia spicata ex hort 5.47 1.92 13.31 1 1 1 1
Diospyros dichrophylla EC 11.65 5.32 217.49 1 1 1 1
Euclea natalensis EC 5.18 4.96 95.91 1 1 1 1 1
Euclea racemosa EC 4.77 4.73 74.75 1 1 1 1 1
Euphorbia mammillaris ex hort 3.18 1.90 6.61 1 1 1
Euphorbia tetragona EC 2.88 2.09 7.45 1 1 1 1 1
Grewia occidentalis ex hort 5.37 3.08 59.37 1 1 1 1 1
Grewia robusta EC 5.34 3.72 40.74 1 1 1 1 1
Gymnosporia capitata 3.33 1.63 3.97 1 1
Hippobromus pauciflorus ex hort 5.43 4.01 60.04 1 1 1
Maytenus heterophylla EC 4.03 2.39 14.8 1 1 1 1 1
Mystroxylon aethiopicum
subsp. aethiopicum MP 11.19 8.32 383.53 1 1 1 1 1
Olea europaea subsp.
africana EC 10.38 7.73 380.39 1 1 1 1 1
Pappea capensis 6.39 5.12 119.51 1 1 1 1
Ptaeroxylon obliquum EC 6.19 1.16 14.14 1 1 1 1
Putterlickia pyracantha EC 5.62 2.72 32.3 1 1 1 1 1
Schotia brachypetala ex hort 5.18 5.18 35.71 1 1 1 1 1
Scutia myrtina EC 13.76 2.38 1 1 1 1 1
Zanthoxoylum capense 4.21 3.14 25.4 1 1
*100 seeds were weighed for Brachylaena elliptica

The cold stratification treatment to determine whether chilling stimulates germination
entailed storing the seeds in a cold room at 5C for two weeks. Germinating seeds in a
nutrient-free medium, such as river sand, was done to determine whether the nutrient
content of soil stimulates germination of Thicket species. Seeds were germinated under
80% shade cloth to investigate the effect of the quantity of light on germination.

70
It was decided to germinate seeds in a glasshouse and not environmental cabinets, as the
glasshouse had temperature regime similar to that in Thicket. Seeds were placed in seed
trays containing a 1:1 mixture of potting soil and filter sand, except for the river sand
treatment. Three replicates of 33 seeds was used, except for Carissa bispinosa, Diospyros
dichrophylla, Euphorbia mammillaris and Zanthoxylum capense, where three replicates
of 20 seeds was used, due to lower seed availability. The seed trays were placed in the
glasshouse in blocks consisting of the same treatment, and trays were rotated in this block
every two weeks. Germination was measured as emergence from the soil. This may
underestimate germination, but is ecologically more useful. The seed trays were checked
for germination every three days for six weeks. This period of time was chosen, as other
processes apart from the specific treatments may cause germination over the experimental
time (Baskin & Baskin 1998). All treatments were not run concurrently, as space was
limited. The experiment ran from May 2008 to January 2009.

Statistical analysis was done using Statistica 8 (Statsoft 2007). Percentage germination
was log transformed, and subjected to single-factor ANOVA. Tukey post-hoc tests were
used to compare significant differences between the treatment means.

3.3 Results
No germination was recorded in Brachylaena ilicifolia, Carissa bispinosa, Euclea
natalensis, Hippobromus pauciflorus, Ptaeroxylon obliquum, Putterlickia pyracantha,
Scutia myrtina and Zanthoxylum capense, as well as the succulents Aloe africana and
Euphorbia tetragona. Only a single seed germinated for all of the treatments for
Gymnosporia capitata (control) and Diospyros dichrophylla (shade), and Olea europaea
subsp. africana only germinated when scarified (Table 11).

Germination of six species was significantly affected by the various treatments; Acacia
karroo (F = 5.53, n = 6, p < 0.05), Schotia brachypetala (F = 4.12,df = 5, p < 0.05),
Cussonia paniculata (F = 4.06, df = 5, p < 0.05), Euclea racemosa (F = 16.73, df = 5, p <
0.001), Maytenus heterophylla (F = 3.41, df = 5, p < 0.05) and Burchellia bubalina (F =
71
10.50, df = 5, p < 0.001) (Table 11). Significant differences between treatments and the
control were identified in only three of these. Hot water (p < 0.05) and nutrient-free sand
(p < 0.01) resulted in increased germination of Acacia karroo, whereas hot water
prevented germination of Cussonia paniculata (p < 0.05). Burchellia bubalina only
germinated in sand and shade, but only shade resulted in significantly greater germination
(p = 0.001).

Acacia karroo germinated the fastest, as germination was recorded on the third day for
the hot water, sand and shade treatments (Fig. 13). Highest germination for the control of
all the species occurred in Schotia brachypetala (47.5% 4.63%) and Pappea capensis
(24.2% 8.02%).

Scarification with sand paper increased germination in Acacia karroo, but resulted in
reduced germination of Cussonia spicata, Euclea racemosa and Maytenus heterophylla
(Fig. 13). Seeds of Acacia karroo were stimulated by immersion in hot water. However,
the same treatment inhibited germination in Schotia brachypetala, Cussonia spicata,
Euclea racemosa and Pappea capensis. The greatest percentage germination for
Euphorbia mammillaris occurred when immersed in hot water, but this was counteracted
by no germination in another replicate of the treatment. Cold stratification had no
significant effect on any of the species.

Germination in sand resulted in improved germination for Acacia karroo (p < 0.01), but
had little effect on the other species (Table 11). Although shade only significantly
increased germination in Burchellia bubalina (p = 0.001), it did the cause an increase in
the rate of germination in Euclea racemosa, Maytenus heterophylla, Schotia
brachypetala, Pappea capensis and B. bubalina (Fig. 13), and did not inhibit
germination in any of the species.

72
0 3 6 9 12 15 18 21 24 27 30 33 36 39 42 45
Time (days)
0
20
40
60
80
100
G
e
r
m
i
n
a
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i
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(
%
)
Con
MS
HW
CS
Sa
Sh
0 3 6 9 12 15 18 21 24 27 30 33 36 39 42 45
Time (days)
0
20
40
60
80
100
G
e
r
m
i
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a
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(
%
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Con
MS
HW
CS
Sa
Sh
0 3 6 9 12 15 18 21 24 27 30 33 36 39 42 45
Time (days)
0
20
40
60
80
100
G
e
r
m
i
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(
%
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Con
MS
HW
Sh
0 3 6 9 12 15 18 21 24 27 30 33 36 39 42 45
Time (days)
0
20
40
60
80
100
G
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(
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Con
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HW
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Sa
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Time (days)
0
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HW
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Sa
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Time (days)
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Time (days)
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Con
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0 3 6 9 12 15 18 21 24 27 30 33 36 39 42 45
Time (days)
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Time (days)
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0 3 6 9 12 15 18 21 24 27 30 33 36 39 42 45
Time (days)
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G
e
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(
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Con
MS
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Sa
Sh
0 3 6 9 12 15 18 21 24 27 30 33 36 39 42 45
Time (days)
0
20
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60
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100
G
e
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(
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Con
MS
HW
Sa
Sh
0 3 6 9 12 15 18 21 24 27 30 33 36 39 42 45
Time (days)
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100
G
e
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i
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i
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(
%
)
Con
MS
HW
Sh

Figure 13 Germination rates of study species. Con () = control,
MS () = mechanical scarification, HW () = hot water, CS ()
= cold stratification, Sa () = sand, Sh () = shade.

Acacia karroo Cussonia
paniculata
Cussonia spicata
Euclea racemosa
Grewia
occidentalis
Grewia robusta
Maytenus
heterophylla
Mystroxylon
aethiopicum
Olea europaea
Schotia
brachypetala
Pappea capensis
Euphorbia
mammillaris
Burchellia
bubalina

0 3 6 9 12 15 18 21 24 27 30 33 36 39 42 45
Time (days)
0
20
40
60
80
100
G
e
r
m
i
n
a
t
i
o
n

(
%
)
Con
MS
HW
CS
Sa
Sh
73
Table 11 Mean (n = 3) germination (% SD) of study species exposed to different treatments. Small letters indicate treatments with significant
differences from the control.
Species F p Control Mech Scar Hot Water Cold Strat Sand Shade
mean SD mean SD mean SD mean SD mean SD mean SD
Acacia karroo 5.53 0.01 6.1 3.03 a 14.1 4.63 a 31.3 11.47 b 12.1 10.93 a 41.4 4.63 b 13.1 4.63 a
Burchellia bubalina 10.50 0.00 0 0 a 0 0 a 0 0 a 0 0 a 1.0 1.75 a 7.1 4.63 b
Cussonia paniculata 4.06 0.02 15.2 3.03 a 12.1 6.06 a 0 0 b 5.1 3.50 a 3.0 5.25 a 7.1 9.74 a
Cussonia spicata 2.58 0.10 10.1 7.63 3.0 3.03 4.0 4.63 20.2 7.63 4.0 4.63
Diospyros dichrophylla 1.00 0.46 0 0 0 0 0 0 0 0 0 0 1.0 1.75
Euclea racemosa 16.73 0.00 9.1 5.25 1.0 1.75 0 0 0 0 9.1 6.06 18.2 8.02
Euphorbia mammillaris 0.58 0.65 1.7 2.89 3.3 2.89 8.3 10.41 1.7 2.89
Grewia occidentalis 1.50 0.26 8.1 7.00 11.1 4.63 4.0 4.63 19.2 3.50 19.2 20.18 12.1 10.50
Grewia robusta 0.85 0.54 2.0 1.75 1.0 1.75 1.0 1.75 0 0 0 0 1.0 1.75
Gymnosporia capitata 1.00 0.44 1.0 1.75 0 0 0 0 0 0
Maytenus heterophylla 3.41 0.04 9.1 3.03 2.0 1.75 4.0 4.63 4.0 4.63 3.0 3.03 20.2 7.00
Mystroxylon aethiopicum 2.86 0.06 2.0 3.50 3.0 0 1.0 1.75 5.1 1.75 0 0 1.0 1.75
Olea europaea subsp. europaea 1.00 0.46 0 0 2.0 3.50 0 0 0 0 0 0 0 0
Pappea capensis 2.82 0.08 24.2 8.02 23.2 9.26 12.1 10.93 25.3 9.74 61.6 8.75
Schotia brachypetala 4.12 0.02 47.5 4.63 46.5 30.35 12.1 8.02 45.5 10.50 17.2 11.47 61.6 11.47

74
3.4 Discussion
Dormancy is more important in unpredictable environments for ensuring germination is
limited to periods of conditions that are beneficial for seedling survival and establishment
(Gutterman 1994, Vandvik & Vange 2003). Thicket is subtropical with a relatively
constant climate, but recurrent drought is common (Kerley et al. 1995), and a high co-
efficient of variation of rainfall is experienced (Cowling 1983). It can be assumed that
most species would then exhibit a cueing mechanism to promote germination when water
availability is high, similar to subtropical Western Australian species (Bell et al. 1995).
As Thicket experiences bimodal rainfall, with peaks occurring in spring and autumn, a
mechanism to enhance germination in spring would be beneficial (Vlok et al 2003).
However, this germination requirement does not require a dormancy mechanism, as seeds
will not germinate if they have not imbibed water. Therefore many species may lack
dormancy.

The wider the range of conditions that a seed is able to germinate in, the smaller its
degree of dormancy (Vleeshouwers et al. 1995). As most woody Thicket species are
widespread (Table 9, Cowling 1984, Vlok et al. 2003) and exposed to a wide range of
environmental conditions (Palmer & Pitman 1972), it could be assumed that many lack
dormancy mechanisms that would limit them to specific microhabitats. Many species are
expected to be quiescent and only require imbibition for germination to occur (Bewley &
Black 1994). The lack of germination in species is contradictory to the low number of
significant effects in species that did germinate.

The largest drawback of this study was not knowing the age of the seeds when they were
received. After-ripening leads to increased germination of many species, but not in
recalcitrant species (Teketay & Granstrm 1997). Loss of viability seemed evident in
Aloe africana, Burchellia bubalina and Ptaeroxylon obliquum seeds, as these three
species had the high germination in the seedling survival experiment (data was not
presented) where seeds were collected from the field. The seeds of Thicket species are
regarded as commonly recalcitrant (Sigwela 2004). Olea europaea subsp. cuspidata does
75
not form a persistent seed bank (Teketay & Granstrm 1995), and a similar situation can
be assumed for Olea europaea subsp. africana, as they occupy similar habitats. Poor
germination has also been observed in Ptaeroxylon obliquum and Euclea undulata in
Addo (Todkill 2001), although no indication of seed age is given by Todkill (2001).
However, Davis (2007) found no loss of viability of Acacia karroo and Grewia robusta
seeds over four months, whereas Azima tetracantha, another typical Thicket species, had
significantly increased germination over time.

The period of germination of 45 days used in this study appears to have been sufficient.
Baskin & Baskin (1998) recommends two weeks for a germination study to preclude any
cues such as warm or cold stratification that may occur during the experiment. However,
a longer period of time was deemed necessary in this study, as emergence from the soil
was the measure of germination, and not radicle emergence. Longer than 45 days would
have resulted in less conclusive data gained about the dormancy mechanism of the
various species, as temperature and light fluctuations occurred in the glasshouse. Thus it
would have been inaccurate to assume that the treatment was the cause of breaking
dormancy.

3.4.1 Seed Coat Induced Dormancy
Mechanical scarification of seeds did not result in significantly increased germination in
any of the species, whereas immersion in hot water enhanced germination of Acacia
karroo (p < 0.01), but inhibited Cussonia paniculata (p < 0.05). The difference between
these two species is that A. karroo has a hard seed that is mammal-dispersed, compared to
the bird-dispersed seeds of C. paniculata that lack a thick seed coat. Dormancy in hard
seeds is broken when the seed coat or other specialised water entry structure is made
permeable (Bewley & Black 1994, Li et al. 1999). This can occur by animal ingestion
(Davis 2007), fire and high temperatures (Jeffrey et al. 1988, Bell et al. 1995, Mucunguzi
& Oryem-Origa 1996), large fluctuations in diurnal temperatures (Vasquez-Yanes &
Orozco-Segovia 1993, Van Klinken & Flack 2005), abrasion by soil particles in
unconsolidated soils (Gutterman 1994), as well as time (Teketay & Granstrm 1997).
76
Species with hard seeds usually occur at the edge of forests and open woodlands, where
they are exposed to the previously mentioned ecological factors (Teketay & Granstrm
1997). This promotes establishment in canopy gaps, where light intensity is greater, a
requirement for the growth of many shade intolerant species. Hall (2002) identified
Acacia karroo and Diospyros dichrophylla, that both have hard seeds, as pioneer or early
successional species in Thicket, indicating that species in Thicket do occur in this
regeneration niche.

Many species in Thicket have relatively thick seed coats (pers. obs.). Thick seed coats are
an adaptation to promote establishment in open areas, as well as to survive the passage
through the gut of an animal or protection against granivory (Sigwela 2004). Species with
hard seeds or thick seed coats may preferentially establish in the open, or on the edge of
Thicket, where there is greater exposure to the outside environment, and larger
temperature fluctuations occur. Establishment in gaps may not be beneficial in Thicket,
especially in drier areas, due to the severe temperature and water stress (Lechmere-Oertel
2003). However, Thicket species of Savanna origin may require canopy gaps for
recruitment (Holmes & Cowling 1993). Nonetheless, thicker seed coats may rather be an
adaptation to endozoochory in Thicket, than assuring seedling germination occurs in
canopy gaps and other open areas. Endozoochory is the preferential mode of dispersal, as
birds spend most their time in the bush, and avoid open interspaces (Bruton 1992,
Alcantara et al. 2000), depositing the seeds into the Thicket clumps, where seedling
establishment is improved (La Cock 1992, Sigwela 2004).

Breaking seed coat imposed dormancy by immersion in hot water is effective in many
species (van Klinken & Flack 2005), including Acacia karroo (Fig. 13). However,
dormancy is not affected by gut passage for A. karroo (Davis 2007) as well as A. erioloba
(Barnes 2001). Shade promoted germination and seedling survival of A. karroo, but
resulted in etiolated seedlings (OConnor 1995). The improved germination of A. karroo
in sand is difficult to explain. A lack of germination from cattle dung is evidence of no
nutrient sensitivity (OConnor 1995), but does not explain the enhanced germination.
Germination is usually greater in dark coloured soils compared to lighter ones, as a result
77
of increased maximum and minimum temperatures (Ludwig & Harper 1958) and this
may have been the case in this study.

The extremely low germination rates of Grewia robusta are similar to those found by
Davis (2007), who compared ingested seeds to uningested seeds. The removal of the
thick seed coat by domestic pigs resulted in improved germination, and improved
germination after passing through elephants (Davis 2007), suggests that seeds of G.
robusta may have coat-imposed dormancy.

It was expected that Schotia brachypetala would have enhanced germination when
exposed to scarification or hot water, as it has a hard seed. However, this was not the
case, with these treatments having no significant effect (Table 11). Germination of S.
brachypetala seeds occurred quickly in all treatments. Not all hard seeds are impermeable
to water (Baskin & Baskin 1998). It can therefore be assumed that S. brachypetala lacks
seed coat-imposed dormancy, even though it has a hard seed. Germination greater than
90% was observed in S. afra without scarification (Todkill 2001). Rather, a hard seed
coat may be a mechanism to protect against granivory (Sigwela 2004).

3.4.2 Embryo Dormancy
Shade resulted in increased germination in a number of species, but only significantly
increased germination in Burchellia bubalina. This species has small seeds with a thin
seed coat and occurs mainly in forest (Coates-Palgrave 2002). It is a shade-tolerant
species that prefers high soil moisture (Chapter 4). B. bubalina is adapted to regenerate
under canopy cover, where the quantity of light is much reduced, compared to the open.
The requirement for shade conditions prevents germination in the open, where poorer
seedling survival would occur (Lieberman & Li 1992). It was observed that seedlings of
many species were highly etiolated in the shade, an indication that other species were
adapted to shade too (pik & Rolfe 2005).

78
Shade has been observed to promote more rapid and complete germination of other
Thicket species, such as Euclea racemosa, Putterlickia pyracantha and Sideroxylon
inerme (Cowling et al. 1997). Low light intensity is an adaptation to ensure seed burial
before germination (Bell et al. 1995). The improved germination of Pappea capensis is in
contrast to its poor growth and photosynthetic performance in shade (Holmes & Cowling
1993). However, P. capensis occurs in a wide range of climatic conditions, which is
reflected in its leaf morphology, as it has very large leaves in high rainfall areas, and
much reduced leaves in inland xeric areas (Coates-Palgrave 2002, pers. obs.). Thus
genetic plasticity allows this species to recruit in many different environments, and it is
one of the dominant seedling species in xeric Thicket (Sigwela 2004).

In species that have embryo dormancy, light is the primary inducer of germination
(Vleeshouwers et al. 1995). Light limits the germination of many Western Australian
species, especially at higher temperatures (Bell et al. 1995). This is a mechanism to
prevent surface germination, as rainfall is strongly seasonal, and soils dry out readily
(Bell et al. 1995). Dormancy is broken by exposure to light in many species (Pons 1991),
but these occur in habitats where water is not limiting. Another observation was that the
soil retained water better underneath the shade cloth compared to the other treatments,
especially the sand treatment. The improved soil moisture retention may have enhanced
germination (OConnor 1995). Germination is very sensitive to soil water, as imbibition
is governed by soil water potential and species respond in different ways (Evans &
Etherington 1990). The failure to germinate at low soil water potentials is a mechanism to
promote successful seedling establishment. Although soil moisture and nitrates do not
affect dormancy, they do strongly influence germination (Vleeshouwers et al. 1995), a
further indication that a specific dormancy mechanism may be lacking.

As Thicket is subtropical, species may not require a season of lower temperatures to
promote germination. This could explain the lack of response to cold stratification.
However, germination in Sideroxylon inerme, a common species in Dune Thicket, is
strongly seasonal, and only occurs in spring (pers. obs.). Sideroxylon inerme may rather
respond to daylength than temperature (Bewley & Black 1994).
79
Two groups of species with embryo dormancy may occur, based on their regeneration
niche. Species whose germination is enhanced by light and large fluctuations in
temperature are likely to regenerate in the open, whereas species that are stimulated by
dark would establish underneath the canopy. It is suspected that it would not be possible
to identify to which group a species belongs just by physical appearance.

Wind-dispersed species, such as Aloe africana, Brachylaena elliptica and Ptaeroxylon
obliquum may not require specific dormancy mechanisms to ensure successful
establishment, as the wind should deposit them into a safe site that is protected from
the environment (Fowler 1988). Nutrient-rich litter may also be blown there (Fowler
1988). Large quantities of seed are also commonly produced, to further ensure successful
regeneration (Westoby et al. 1996).

Of the three succulent species in this study, only Euphorbia mammillaris germinated. No
treatment resulted in significantly increased germination, but immersion in hot water
resulted in more rapid germination. This may indicate a seed coat-imposed dormancy, but
germination was too low to make any conclusive deduction.

3.4.3 Conclusion
Knowledge of the seed biology is essential to understanding community processes such
as plant establishment, succession, and natural regeneration (Vasquez-Yanes & Orozco-
Segovia 1993). It is also critical for the rehabilitation of ecosystems (Bell et al. 1993).
This study furthers this understanding for Thicket, and hopefully will contribute to our
understanding of regeneration.

Many Thicket species did not germinate over a period of 45 days. Low percentage
germination was recorded in many species, and no germination occurred in ten of the 25
species. This prevents any clear picture from emerging about the dominant dormancy
mechanisms that occur in woody and succulent canopy species in Thicket. It is likely that
most species are quiescent and lack dormancy. Thus it would not be acceptable to accept
80
or reject the hypotheses in this study, due to the limited germination success. However,
immersion in hot water did result in improved germination in one species with a thick
seed coat, but inhibited germination in a thin seed-coated species. This finding, together
with the fact that shade only resulted in significantly improved germination in a species
with a thin seed coat, allows both hypotheses to be accepted tentatively.

It can be predicted that species with hard seeds (Acacia karroo, Diospyros dichrophylla,
Schotia brachypetala) would establish preferentially in the open, or on the edge of
Thicket clumps. Species with thick seed coats (Olea europaea subsp. africana) would
mostly regenerate underneath the canopy, as that is where they are dispersed. These are
usually large seeds that contain substantial reserves, resulting in improved seedling
establishment and survival. Species that lack thick seed coats could be divided into two
groups: Pappea capensis, Burchellia bubalina and Euclea racemosa are bird-dispersed
species with improved germination in shade, and would be expected to establish
underneath the canopy. Other species that are inhibited by shade and respond to
fluctuations in temperature or light may regenerate in the open. However, this group was
not assessed in this study, but may include species of Lycium.

81
Chapter 4 Response of seedlings to drought and shade
4.1 Introduction
The process of the regeneration of a community is completed with the establishment of
seedlings, which is usually the most important phase in terms of survival (Harper 1977,
Rey et al. 2000, Fenner & Thompson 2005). However, this establishment is limited by a
number of causes of seedling mortality, including competition for light, space and
nutrients from other seedlings as well as existing vegetation, herbivory, physical damage
as a result of branch falls, and lack of moisture (Fenner & Thompson 2005). In the
Knysna Forest, high levels of emergence of seedlings occur in Apodytes dimidiata, a
species common to both Forest and mesic Thicket, but numbers are quickly thinned out
by fungal attack and herbivory by rodents and antelope (Phillips 1927). Seedlings that
survive to the sapling stage are further targeted by antelope, resulting in few individuals
reaching adulthood. Any seedlings that are dispersed into open sites die immediately
from sun scorch (Phillips 1927).

4.1.1 Seedling response to drought and shade
The establishment and persistence of a species is dependent on its ability to perceive and
adapt to changes in their environment (Sack & Grubb 2002). As the adaptive plasticity of
plants promote environmental tolerance, differences in the plasticity patterns of
individual species may explain differences in their ecological breadth (Griffith & Sultan
2005). An understanding of the response of seedlings to the effects of shade and drought
is essential for explaining species niche differentiation, and will allow a prediction of the
survival and growth of seedlings in different microsites (Sack & Grubb 2002).

Plants differ widely in their capacity to survive drought conditions and can be classified
according to the dominant mechanisms employed. Drought avoidance occurs in species
that occur in regions that experience well-defined dry seasons, where species are able to
complete the growing phase of their life cycle during the wet season (Kramer &
Kozlowski 1979, Gullo & Salleo 1987). Species that can tolerate drought may be divided
82
into two groups. Woody species are classified as desiccation tolerators as they can endure
extreme dehydration of protoplasm and low relative water content (RWC), or avoid low
water potential (
1
) without irreversible injury (Kozlowski & Pallardy 2002).
Desiccation avoider species have a number of mechanisms that allow survival, including
leaf adaptations such as leaf shedding; small or few leaves; leaf pubescence; small, few
and sunken stomata; rapid stomatal closure during drought; abundant leaf waxes and
strong development of palisade mesophyll (Gullo & Salleo 1987, Burslem et al. 1996,
Sandquist & Ehleringer 1997, Gindaba et al. 2004). Stems are adapted to drought by
transferring photosynthetic processes to stems and twigs, as well as having a low
resistance to water flow in vascular tissues (Meinzer 2002), whereas roots adaptations
include extensive root growth, increased root:shoot ratios, increased lateral root growth as
well as high root regenerating potential after transplanting (Burslem et al. 1996,
Kozlowski & Pallardy 2002).

Drought and desiccation is the main cause of seedling mortality in Mediterranean
shrublands (Rey et al. 2000, Arrieta & Surez 2006), Savanna (OConnor 1995, Barnes
2001) and dry tropical Forests (McLaren & McDonald 2003, Asbjornsen et al. 2004), and
prevents the establishment of trees in arid areas (Kozlowski & Pallardi 2002). Seedling
establishment of Acacia karroo in years that experience less than 500 mm of rainfall is
poor, and establishment is limited to years of average rainfall in the mesic grasslands of
the Eastern Cape (OConnor 1995). A similar pattern may occur in Thicket, where the
seedlings of many species can only establish during years when rainfall is above a certain
amount. These rainfall events could be exceptional in Thicket types that experience lower
rainfall, causing seedling establishment and regeneration to be episodic (Holmes &
Cowling 1993).

Plasticity of plants can be identified in their response to shade. Plants are able to perceive
both the quality and quantity of light using phytochrome, with different species-specific
responses elicited by changes in the light environment (Holmes et al. 1983, Griffith &
Sultan 2005). Species are able to adapt to various light intensities by changes in the size
and efficiency of their photosynthetic system (Loach 1967), as well as morphological
83
adaptations (Loach 1970). Based on these changes, species can be divided into shade
tolerant (shade) and shade intolerant (sun) species (Holmes et al. 1983, Lee & Graham
1986). Species with phenological plasticity generally respond to shade by increasing their
light-harvesting traits (Griffith & Sultan 2005). This includes increasing the area of
individual leaves, increasing the area of leaves displayed to light (leaf angle) and less
branching to maximise height (Canham 1988, Holmes & Cowling 1993). Photosynthesis
in shade tolerant plants is saturated at a lower photosynthetic flux density (PFD), but has
a low light compensation point, the value of irradiance at which photosynthesis exactly
equals respiration (Loach 1967, Lee & Graham 1986, Kitajima 1994). The photosynthetic
flux density at which the low light compensation point occurs for shade plants is around
20 mol m
-2
s
-1
and 80 mol m
-2
s
-1
for sun plants (pik & Rolfe 2005), enabling shade
plants to survive in light conditions that cannot support sun plants, whereas bright light
will result in shade plants being outcompeted as they are less efficient at
photosynthesising at high irradiances, as well as being more susceptible to photochemical
damage, compared to sun plants (Pags et al. 2003). However many species are able to
adapt to different light levels, as they are able to produce sun leaves outside of the
canopy, and shade leaves within it (Bjrkman 1981). These leaves operate in a similar
manner to sun and shade plants (Bjrkman 1981).

A summary of possible plant responses to the combined effects of drought and shade are
presented in Sack & Grubb (2002). Shade is predicted to have a stronger impact due to
lower investment in root biomass (Loach 1970), or a weaker impact as a result of
facilitation (Sigwela 2003) or light-limited growth, or equal and proportional impact on
seedlings growing in deep shade compared to individuals growing in moderate shade or
in the open (Sack & Grubb 2002).

4.1.2 Seedling growth and survival in Thicket
The transitional nature of Thicket is reflected in its species response to shade and
drought. In xeric areas of Thicket, species need to be able to tolerate extreme
temperatures, as well as extended periods of drought. In mesic areas, greater competition
84
and higher canopy cover results in species requiring fast growth rates, or tolerance of
deep shade conditions. As Thicket has tropical affinities (Cowling 1983), the tropical
forest model could best describe its regeneration patterns (Holmes & Cowling 1993).
Shade intolerant species benefit from canopy gaps, as they are associated with higher
nutrient and light conditions in mesic areas (Pags et al. 2003). In xeric regions, this
relationship does not hold (Pags et al. 2003).

There is a distinct lack of research done on the growth dynamics of Thicket plant species.
Holmes & Cowling (1993) described three guilds of species, according to their response
to shade. Cassine peragua and Sideroxylon inerme were adapted to grow in deep shade,
and are similar to primary tropical species as they do not require openings in the canopy
for seedling establishment. Rhus glauca is a typical pioneer species with a high growth
rate and would greatly benefit from canopy gaps, but has a wide irradiance tolerance and
may be able to establish in shade. Pappea capensis and Schotia afra differ from the
tropical forest model as they have low growth rates but are able to photosynthesise at
high rates in full sunlight. They are adapted to xeric conditions and naturally occur in
open savanna vegetation as well as Thicket, and would require canopy gaps as they
accumulate biomass slowly in shade.

However, few seedlings are found outside Thicket clumps, as extreme environmental
conditions prevent most woody species from establishing. Although competition can
explain the spatial distribution of seedlings, especially shade tolerant species (Pags et al.
2003), this is unlikely in Thicket. The only seedlings found in Xeric Succulent Thicket
were Pappea capensis and Euclea undulata, species adapted to drought conditions
(Lechmere-Oertel 2003; Sigwela 2004). Plant species in Thicket are exposed to a wide
variety of environmental conditions, from semi-arid with rainfall as low as 200 mm and
temperatures peaking at 50C, to subtropical conditions with rainfall as high as 1 050 mm
(Vlok et al. 2003). Rainfall is bimodal with peaks in spring and autumn. However,
rainfall is not reliable as monthly coefficients of variation are high and prolonged
droughts occur frequently (Cowling 1984). Soil in open areas is compact and may
prevent the burial and emergence of seedlings (Todkill 2001). Herbivory is the major
85
cause of disturbance in Thicket, and may be responsible for high levels of seedling
mortality (Kerley et al. 1995). Drought is not regarded as an important source of
defoliation and disturbance, as it does not result in a large change in biomass (Kerley et
al. 1995). Many species have wide distributions in Thicket and are able to tolerate large
ranges of environmental conditions (Coates-Palgrave 2002).

The restoration of Thicket over large areas is necessary (Hoffman 1987, Stuart-Hill
1991), and it is therefore vital to understand the growth and survival responses of
indigenous species to dominant environmental factors (De Ridder 2006). This would be
instrumental in the selection of nurse species that need to be able to survive harsh
conditions without taking an overly extended period of time to grow (La Cock 1992). The
various regeneration niches required by Thicket species also need to be recognised to
fully restore the landscape to its previous composition (Holmes & Cowling 1993, Hall
2000, De Ridder 2006).

The aims of this chapter is to investigate the responses of Thicket seedlings to shade and
drought, to determine if it is one of these factors that prevent seedlings from establishing
in openings in the Thicket. Seedling survival is reported to be highest at the edge of
bushclumps where environmental conditions are intermediate (De Ridder 2006).

The objectives of this study are to:
1. determine which environmental factor is dominant in preventing seedling
establishment in Thicket;
2. describe the responses of the various species to shade and drought.

Hypotheses:
H1: Species with a xeric distribution have higher growth rates in mid-light environment
compared to species with a mesic distribution having highest growth rates in low light.
H2: Species with a xeric distribution would be able to continue growing at drought levels
of moisture, compared to species with mesic distribution.

86
4.2.1 Study species
The initial selection of species was based on Everards (1987) characteristic species of his
various Thicket types. However, due to low rates of germination and restrictions in terms
of glasshouse space, four species were used in this experiment; Aloe africana Mill., Rhus
dentata Thunb., Burchellia bubalina (L.f.) Sim and Ptaeroxylon obliquum (Thunb.)
Radlk.

Aloe africana is an arborescent succulent of the family Asphodelaceae (Coates-Palgrave
2002). It can grow up to 4 m and does not branch. It occurs within dense Thicket and
maintains its old dry leaves around the stem. It is limited to the south-western part of the
Eastern Cape and is commonly known as the Uitenhage aloe. It has spreading leaves that
are dull green in colour and have reddish teeth at the edge as well as a few on the upper
and lower surface of the leaves. The leaves start horizontal, curving down as they get
older. Inflorescences consist of branched heads, with 2 or 3 heads per leaf rosette.
Flowers are bright yellow-orange, 40-60 cm long and curve upwards. Flowering occurs in
July to September.

Ptaeroxylon obliquum, otherwise known as sneezewood, is a shrub to tall tree up to 20 m
in height of the family Ptaeroxylaceae (Coates-Palgrave 2002). It is not limited to the
Eastern Cape, stretching from the western half of the Eastern Cape up to Zimbabwe along
the eastern seaboard. It occurs in lowland Savanna and Thicket to mistbelt Forest. It has
pale grey bark that is longitudinally fissured and prone to flaking. Leaves are deciduous,
paripinnate, opposite and crowded at the ends of the branchlets. The rachis is slightly
winged, 3-7 pairs of opposite leaflets occur which are obliquely rectangular and dark to
blue-green in colour. Flowers are white to pale yellow, individual trees are monecious.
Fruit are oblong, apically-notched capsules that become reddish-brown when mature, and
split into two valves to release a pair of winged seeds. Flowering occurs August to
December, and fruiting December to February.

87
Rhus dentata, a member of the Anacardiaceae family, is a shrub to small tree up to 5 m
tall, occurring from the western half of the Eastern Cape into KwaZulu-Natal and the
highveld of the Free State, Mpumalanga, Limpopo, Gauteng and North-West Provinces
(Coates-Palgrave 2002). It grows in a variety of habitats, from rocky hillsides, Thornveld
to Bushveld, at the margins of evergreen Forest in riverine vegetation. Its bark is greyish
brown, smooth or striated, and its compound leaves consist of three leaflets, obovate to
widely elliptic, with the lateral leaflets half the size as the terminal one. Leaves are thinly
textured, pale to dark green above, paler with prominent midrib below, with or without
hairs. Leaves turn dull yellow to orange-red before falling off. Small flowers are yellow
in colour, occurring in axillary heads up to 14 cm long. Flowering occurs from November
to March.

Burchellia bubalina is a shrub to small tree up to 5 m in height, a member of the
Rubiaceae family (Coates-Palgrave 2002). It is found in evergreen Forest, at Forest
margins, as well as in Grassland on rocky outcrops, occurring in a contiguous strip along
the coastal plain from Cape Town up to the Escarpment of Mpumalanga and Limpopo.
The bark is greyish brown and mottled, and leaves are opposite, broadly ovate, usually
about 10-13 cm long. Leaves have a softly to thickly leathery texture, glossy dark green
above, hairless or finely hairy and dull green below. The margin is entire and the petiole
is thickset. Stipules are leathery, toothed, falling early to leave a conspicuous ridge.
Flowers are scarlet to orange, urn-shaped, occurring in terminal, shortly branching heads
that consist of four to eight flowers. Fruit are urn-shaped and leathery, with persistent
calyx lobes. Seeds are very small and disk-shaped. Flowering occurs from September to
November, and fruiting from November onwards for months.

4.2.2 Germination of seeds
Seed was purchased from nurseries in December 2006. They were germinated on paper
towel in trays covered with plastic wrap under laboratory conditions. After the radicle
had emerged, seedlings were transplanted to seedling trays filled with an equal mixture of
potting soil and filter sand. The seedling trays were placed in the greenhouse and watered
88
every second day, until the experiment began in August 2008. Seedlings were
progressively transplanted into pots when they outgrew the seedling trays, from August
2007 to March 2008. Kelpak was applied to seedlings before the commencement of the
experiment. At the beginning of the experiment, seedlings were between 12 and 18
months old.

4.2.3 Experimental design
To test the response of seedlings of the selected species to different treatments of drought
and light intensity, a split-plot experiment design was used. The experimental design was
a latin square, with four levels of shading: full ambient light (no shade), 40%, 60% and
80% shade; and three levels of water treatments: 85%, 50% and 10% of field capacity.
The different light intensities were reached using shade enclosures with various
percentages light penetration. The seedlings were randomly placed on four tables. Each
table consisted of a different shade treatment, with the table divided into the three water
treatments. Tables, water treatments and seedlings were rotated on a monthly basis. Five
seedlings were used for each water treatment, making up 15 seedlings in each shade
treatment and a total of 60 seedlings for each species.

The method to determine the field capacity of the soil followed Du Preez (1987). Field
capacity of the soil was measured by placing 5 replicates of 500 g of soil in pots with
filter paper at the bottom. The soil was saturated, allowed to drain for 24 hours and wet
weight was measured. The soil was oven dried at 70C for 48 hours and dry weight was
measured. The field capacity of the soil was expressed as the amount of water in grams
per gram dry mass of soil, multiplied by 100 to convert to percentage.

Initial seedling size measurements were taken on 15 and 16 August, 2008. Only non-
destructive measurements were taken: the number of leaves was counted, stem height was
measured using a piece of string and a tape measure, and stem diameter were measured
with a digital vernier calliper. Leaf length was also measured as a representation of leaf
size. Care was taken to measure stem diameter at the same location on the stem of the
89
individual seedlings. Post-treatment measurements were taken three months later on 17
and 18 November, 2008. Seedlings were watered every second or third day, depending on
the weather.

Statistica 8 (Statsoft 2007) was used for the statistical analysis. Main effects ANOVA
was used to determine overall effects, with Kruskall-Wallis analysis of ranks used to
identify significant differences for shade and water treatments within each species.

4.3 Results
There was a significant species and water effect for all growth variables, but stem and
leaf lengths were the only variables that exhibited a significant shade effect (Table 12).

4.3.1 Effect of shade
Stem diameter: Stem diameter was not significantly affected by shade for all four species
(Fig. 14a). A clear trend of decreasing stem diameter with increased shade is evident in
Aloe africana as well as Burchellia bubalina with the exception of ambient light, where
mortality resulted in lower values at the end of the study. A similar pattern is evident in
P. obliquum with the exception of 60% shade. The amount of light received by R. dentata
resulted in significant changes in the stem diameter (H = 8.85, n = 60, p = 0.031),
although no individual treatments were significantly different from each other. R. dentata
showed greatest diameter growth under the 80% shade treatment.

Table 12 F-values for measured responses of seedlings of the four study species to various shade and
drought treamtments.
Factor n Leaf number Leaf length Stem length Stem diameter
Species 4 2.7* - 119.9*** 10.2***
Shade 4 2.0 3.6* 6.4*** 0.4
Water 3 5.3** 4.7* 3.6* 11.1***

90
Stem height: Stem height was significantly greater under 80% shade compared to open
for A. africana (H = 21.88, n = 60, p < 0.001) and R. dentata, with a clear trend of
increasing stem height with increased shade (Fig.14b). Burchellia bubalina and P.
obliquum had greatest stem height under 60% shade. Seedlings of B. bubalina growing
under 60% were significantly taller than those under 40% shade (p < 0.001). Rhus
dentata stem height (H = 14.34, n = 60, p = 0.003), Seedlings under 80% shade were
significantly taller than those in the open (p=0.008) and under 40% shade (p=0.031).

Leaf number: An increase in shade only caused a significant increase in the number of
leaves in P. obliquum (H = 9.27, n = 60, p = 0.026), with seedlings under 60% shade
having significantly more leaves than under ambient light (p = 0.033). The greatest leaf
loss was under the ambient light treatment, except R. dentata which shed most leaves at
40% (Fig. 14c). Maximum leaf gain was recorded under 60% shade for R. dentata and P.
obliquum. Burchellia bubalina did not show leaf gain in any of the shade treatments,
whereas the maximum increase in the number of leaves for A. africana was 3.

Leaf length: Shade had a significant effect on leaf length of A. africana (H = 8.41, n =
120, p = 0.038), B. bubalina (H = 22.59, n = 98, p < 0.001) and R. dentata (H = 20.54, n
= 102, p < 0.001) with leaves under 60% shade significantly longer than those in 40%
shade (p = 0.023, p < 0.001, p < 0.001 respectively) (Fig. 14d). Leaves of R. dentata
growing under 80% shade were significantly longer than seedlings growing in ambient
light (p = 0.026). Maximum leaf length occurred in all species under 60% shade. In R.
dentata and P. obliquum there was a clear increase in leaf length from ambient light to a
maximum under 60% shade. A similar trend was evident in A. africana and B. bubalina
except that the shortest leaves occurred under 40% shade.

4.3.2 Effect of water
Stem diameter: The amount of water received had a significant effect on stem diameter of
Burchellia bubalina (H = 15.01, n = 60, p < 0.001), Ptaeroxylon obliquum (H = 7.19, n =

91

A. africana R. dentata B. bubalina P. obliquum
0
0.4
0.8
1.2
1.6
2
S
t
e
m

d
i
a
m
e
t
e
r

(
m
m
)
0%
40%
60%
80%
0
10
20
30
40
50
S
t
e
m

h
e
i
g
h
t

(
m
m
)
0%
40%
60%
80%

a) b)
-20
-10
0
10
20
L
e
a
f

n
u
m
b
e
r
0%
40%
60%
80%
0
40
80
120
160
L
e
a
f

l
e
n
g
t
h

(
m
m
)
0%
40%
60%
80%

c) d)
Figure 14 Mean growth responses (a = stem diameter, b = stem height, c= leaf number, d = leaf
length) of Thicket species to various shade treatments in a glasshouse. Lower case letters indicate
significant differences within species. Bars indicate standard error of the mean.

60, p = 0.027), Rhus dentata (H = 11.71, n = 60, p = 0.003) and Aloe africana (H = 6.34,
n = 60, p = 0.042). Stem diameter was significant greatest at 85% field capacity for all
species, except A. africana which peaked at 50% (Fig. 15a). However, no individual
treatments of A. africana were significantly different from each other. Seedlings of
Burchellia bubalina, Ptaeroxylon obliquum and Rhus dentata receiving 85% field
a
a
b
ab
ab
b
a
a
a
ab
ab
b
ab
ab
a
b
bc
a
ab
c
b
a
a
ab
92
capacity had significantly greater diameter than those at 10% (p < 0.001, p = 0.033, p =
0.002 respectively).

Stem height: The amount of water received had a significant effect on stem height of
Burchellia bubalina (H = 18.64, n = 60, p < 0.001). Increase in height was greatest for all
species at 50%, except for B. bubalina which peaked at 85% (Fig. 15b). Seedlings of
Burchellia bubalina growing in 85% field capacity were significantly taller (p < 0.001)
than those seedlings receiving 10%.

Leaf number: The amount of water received caused a significant change in both the leaf
number (H = 14.32, n = 59, p < 0.001) of P. obliquum as seedlings receiving 10% field
capacity lost significantly more leaves than 85% field capacity (p = 0.003) (Fig. 15c).
Increase in leaf number was greatest for all species at 85% field capacity, except for P.
obliquum that peaked at 50%. The number of leaves of B. bubalina differed significantly
with the field capacity of the soil (H = 10.76, n = 60, p = 0.005) as seedlings receiving
50% (p = 0.028) and 85% (p = 0.008) lost significantly fewer leaves seedlings grown in
10%. The number of leaves (H = 6.80, n = 60, p = 0.033) of R. dentata were significantly
affected by the amount of water received. Seedlings of R. dentata receiving 85% field
capacity had both significantly more leaves (p = 0.029) than the seedlings receiving 10%
field capacity.

Leaf length: Water also resulted in significant changes in leaf length of Burchellia
bubalina (H = 9.09, n = 98, p = 0.010), as leaves of seedlings receiving 85% field
capacity were significantly shorter than those under 10% field capacity (p = 0.008) (Fig.
15d). Leaf length in Aloe africana was also affected by the field capacity of the soil (H =
9.19, n = 120, p = 0.010), with leaves of seedlings growing in 50% field capacity
significantly longer than seedlings in 85% ( p = 0.007). Leaf length was greatest at 50%
for all species, with the exception of P. obliquum which reached maximum leaf length at
85% field capacity.

93
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c) d)
Figure 15 Mean growth responses (a= stem diameter, b = stem height, c = leaf number, d = leaf
length) of Thicket species to various water treatments in a glasshouse. Lower case letters indicate
significant differences within species. Bars indicate standard error of the mean.

4.3.3 Effect of species
Rhus dentata had the greatest increase in stem height and stem diameter of the three
woody species in all individual water and shade treatments. Burchellia bubalina showed
second greatest increase in height, whereas Ptaeroxylon obliquum had second greatest
a
b
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94
diameter growth. The succulent Aloe africana exhibited greater increase in stem diameter
compared to stem height, relative to the woody species.

4.4 Discussion
The light environment underneath the shade-cloth enclosure is not identical to that
experienced underneath the canopy of a Thicket clump, as light flecks are excluded
(Henry & Aarsen 1997, Naumberg & Ellsworth 2000), and there is a smaller alteration of
the red:far red light ratio (Griffith & Sultan 2005). However, many shade tolerant species
do not respond to reduced red: far red light ratio (Holmes et al. 1983) as little benefit is
derived from stem elongation in species under tall canopies (Van Riensberg & Van
Tienderen 1997). Notwithstanding, this experiment still allowed the investigation of the
relative responses to shade of the four species in this study, inferring their preferred
conditions for establishment, as well as their ability to tolerate drought.

It is also important to state that the soil was not kept at a certain percent field capacity,
but the measure of field capacity was used to determine how much water to give to the
various drought treatments. Higher evapotranspiration in unshaded conditions would
have caused soil to have a lower percentage of field capacity for all water treatments.
This effect would have been removed as results were expressed as pooled values
separately for the drought and shade treatments (i.e. all seedlings grown in 80% shade,
disregarding how much water they received, were pooled to identify effect of 80% shade
on growth measurements, and vice versa). A decrease in replicates would be the only
remaining consequence of mortality.

4.4.1 Overall growth responses
Rhus dentata grew considerably faster, both in terms of height and stem diameter,
compared to the other species, as well as having the greatest relative increase in stem
height with increasing shade. Burchellia bubalina and Ptaeroxylon obliquum had lower
growth responses, and Aloe africana was consistently the slowest growing species in
terms of leaf number and stem height.
95

Leaf shedding occurred in all four species during this study. A possible explanation is
that Ptaeroxylon obliquum is a deciduous species and Rhus dentata is regarded as semi-
deciduous (Coates-Palgrave 2002). However, only seedlings of woody species receiving
10% field capacity completely lost all their leaves. This may have been a response to the
change in conditions by acclimating to the lower soil moisture (Bjrkman 1981).

4.4.2 Response to shade
All four species were able to increase stem height and diameter under 80% shade. This
indicates a degree of shade tolerance in all four species. The dominant response to shade
was an increase in the length of leaves in all species, except Ptaeroxylon obliquum. An
increase in leaf size or leaf surface area in shade is a well-documented response to low
light (Loach 1970, Holmes et al. 1983, Canham 1988, Holmes & Cowling 1993, Kitajima
1994, Henry & Aarsen 1997, van Hinsberg & van Tienderen 1997, Sack & Grubb 2002).
Ptaeroxylon obliquum may employ a different strategy of increasing the number of
leaves, rather than leaf size, as shade resulted in a significant increase in leaf number for
this species. Both strategies would result in a greater leaf surface area exposed to light.

No clear pattern was evident for the production of leaves in response to different light
intensities. Although not identified in the data, it was observed that woody seedlings in
the shade treatments produced fewer but larger leaves than seedlings in the open. Plants
usually produce more leaves in full sunlight compared to under canopies (Holmes &
Cowling 1993, Lin et al. 2002). Rhus dentata produced the most leaves, followed by
Ptaeroxylon obliquum. Burchellia bubalina produced a pair of leaves at a time, whereas
A. africana produced a single leaf each time. An individual R. dentata seedling growing
unshaded at 10% field capacity, completely altered its leaf morphology by changing from
a few, medium-sized leaves, to very many tiny leaves. However, only a single R. dentata
responded in such an extreme manner, indicating high levels of morphological plasticity
within the species.

96
A shift away from apical dominance in the three woody species was observed in the
unshaded treatment. Apical dominance occurs in species that respond to shade by
elongating the stem (Henry & Aarsen 1997). A highly branched and bushy architecture
in the open maximises surface leaf area and resultant light interception (Canham 1988).
Another explanation is that leaf shedding resulted in a shift from the dominance of the
apical meristem to the axillary meristems.

Shade did not result in significant changes in the stem height and diameter of B. bubalina
and P. obliquum, classifying these species as shade tolerant. Shade tolerant species are
able to maintain a similar pattern of growth and metabolism over a wide range of light
intensities, by conserving their reserves (Loach 1970, Sack & Grubb 2002). All species in
this study are canopy tree species that will eventually experience full sun, and are thus
facultatively adapted to shade, and not shade obligate species (Kitajima 1994). The
regularly observed decrease in stem diameter in shaded environments (Holmes et al.
1983, Canham 1988, Holmes & Cowling 1993) was not observed in this study, although
the smallest increase in stem diameter did occur under the greatest shade treatment for all
species with the exception of Rhus dentata. The large variability between seedlings of the
same species combined with only five seedlings being used per treatment, makes it
difficult to show significant differences in stem diameter. The seedlings of the three
woody species growing in the unshaded 10% field capacity treatment experienced greater
mortality, reducing overall diameter growth over the period of study, compared to the
other treatments. This would have resulted in smaller difference between the treatments
for 10%.

Rhus dentata seedlings grew tall under low light conditions. Although Loach (1970)
describes this as a form of shade intolerance, mentioning that intolerant species are the
most plastic, this strategy is better known as shade avoidance, whereby plants experience
a rapid increase in stem height as a mechanism to escape the vegetation canopy (Henry &
Aarsen 1997). Not just shade intolerant, but all light-demanding species exhibit a greater
phonological plasticity to the light environment (Kitajima 1994). Increases in the length
of petioles and internodes, as observed in R. dentata, are further indications of shade
97
avoidance (Henry & Aarsen 1997). Much smaller but significant stem growth occurred in
A. africana, thus classifying it as a shade avoider as well. A. africana prefers the dense
bush of Thicket to open areas (Coates-Palgrave 2002), but regularly emerges from the
canopy (Vlok et al. 2003). This succulent species would not be intolerant to shade, as it is
one of the few species whose seedlings are commonly found underneath the canopy of
Thicket (Midgley & Cowling 1993).

4.4.3 Response to drought
The number of leaves produced by woody plants in this study was a function of water
supply, as leaf production was greatest for all three woody species at 85% field capacity.
In contrast, leaf production of Aloe africana was not affected by water supply. A decrease
in leaf number and leaf area has been observed for tropical seedlings as a result of
infrequent watering (Burslem et al. 1996). Leaf loss has been shown to be a drought
tolerance mechanism in a desert shrub (Sandquist & Ehleringer 1997). Leaf shedding
allows the plant to quickly decrease the evapotranspiration surface, limiting the loss of
water (Burslem et al. 1996).

The ability of Thicket species to resprout was evident in this study. Seedlings of woody
species in treatments receiving 10% and less commonly 50% field capacity shed all their
leaves and resprouted new ones. This continued until the beginning of November when a
large number of first the Burchellia bubalina seedlings, followed by Rhus dentata and
finally Ptaeroxylon obliquum seedlings died. However, watering continued throughout
the study due to their resprouting ability. Mortality of the three species began in the
seedlings experiencing full sunlight, but had occurred in all light treatments by the middle
of November when the final measurements were taken. The drought tolerance of P.
obliquum was evident as it was the only woody species to remain alive in the unshaded
10% field capacity treatment a month after the experiment was terminated, but the
various watering treatments had persisted.

98
Rhus dentata changed its leaf morphology under low water treatments in the unshaded
treatment from a few medium-sized leaves to many small leaves, causing a large increase
in leaf number. As new leaves are assumed to be more adaptable than older leaves (Loach
1970), species with a high leaf production are able to take full advantage of changed
environmental conditions. This plasticity allows R. dentata to survive in a broad range of
environments (Holmes & Cowling 1993, Griffith & Sultan 2005). Both R. dentata and B.
bubalina developed leaf pubescence in the unshaded treatment. Pubescence, common in
drought tolerant species, reduces the absorption of solar radiation, as well as reducing the
heat load of the leaf (Sandquist & Ehleringer 1997). Although these species cannot be
described as drought tolerant, they do have adaptations that contribute to improved
survival in dry habitats.

All species, with the exception of A. africana, increased their stem diameter in response
to improved soil water conditions, with maximum diameter growth in seedlings receiving
85% field capacity. The largest relative increase was seen in B. bubalina. The same
pattern of increasing stem diameter with increased water was not reflected in stem height,
as height growth was only significant in B. bubalina. The three other species reached
maximum height at 50% field capacity. As drought tolerance can be defined as the ability
to maintain growth at a lower soil moisture (Sack & Grubb 2002), B. bubalina could be
described as drought intolerant. It has the largest leaves of the woody species, as well as
the largest observed internode lengths (not measured) for seedlings at 85% field capacity.
This is similar to tropical seedlings that experienced reduced growth as a result of
drought (Burslem et al. 1996). The drought tolerance of the four species accurately
predicts their distribution in Thicket, especially as the three woody species reach their
arid end of their distribution in Thicket (Coates Palgrave 2002). Burchellia bubalina is
the most mesic-adapted species, and limited to Mesic Kaffrarian Thicket, whereas Rhus
dentata stretches into Xeric Kaffrarian Thicket (Everard 1987). Ptaeroxylon obliquum
occurs throughout the biome, whereas Aloe africana is limited to Mesic Succulent
Thicket.

99
Drought tolerance was evident in A. africana and P. obliquum, as both species had slow
growth rates, and increased stem diameter as opposed to height, relative to the other
species. Xeromorphic or drought tolerant species have slower rates of growth (pik &
Rolfe 2005). For seedlings growing in shade, durability of the seedling body is more
important than net carbon gain (Kitajima 1994). This would be especially true in Thicket,
as herbivores of all sizes would have a large effect on seedlings, even those growing
under the relative safety of the canopy (De Ridder 2006). Most succulent species have a
low surface:volume ratio, resulting in reduced levels of CO
2
diffusion into the
photosynthetic tissues. As only the outer layers of cells of the photosynthetic organs are
well illuminated, photosynthesis is limited to these layers. The ratio of photosynthetic
tissue to total mass is low, and thus the net assimilation rate in slow with consequent slow
growth (pik & Rolfe 2005). Increased levels of resource investment in roots and
consequent increased root:shoot ratio, as well as in supportive tissues, causes fewer
resources to be available for stem growth. Higher investment in supportive tissue allows
plants to resist cavitation and wilting, allowing species to survive in semi-arid and arid
environments (Meinzer 2003). However, despite environmental influences, species with a
broad range of wood density co-exist in both humid and arid environments, pointing to
genetic control (Meinzer 2003).

4.4.4 Implications for seedling survival and regeneration in different microhabitats
The distribution and niche of species is dependent on the range of environmental
conditions in which it can establish (Gurevitch et al. 2006). Physiological and
morphological plasticity, especially important in immobile plants, allows species to
tolerate a wider range of conditions, increasing their distribution and fitness (Griffith &
Sultan 2005). As all four species in this study can grow in shade, the low-light
microhabitat underneath the Thicket canopy would not prevent establishment. It is likely
that establishment of Burchellia bubalina would be limited to underneath the canopy,
except in areas where rainfall is sufficient to prevent seedling survival. Rhus dentata
would preferentially establish at the edge of Thicket where it would be able to escape the
canopy relatively quickly. Ptaeroxylon obliquum could establish in the open or under the
100
canopy, but survival will be greater at the edge of Thicket as well. Aloe africana is absent
from Kaffrarian Thicket (Everard 1987). This could be a result of increased fire in the
east (Vlok et al. 2003), greater competition from faster growing species, or the inability
to escape the higher canopy of Mesic Kaffrarian Thicket.

Rhus dentata, similar to R. glauca (Holmes & Cowling 1993), can be described as a
pioneer species, although limited to mesic areas. Small trees and shrubs experience high
variability in light intensity under early successional canopies (Henry & Aarsen 1997),
selecting for highly plastic species that adapt quickly to changing light conditions. The
edge of Thicket clumps would have a similar light environment to an early successional
canopy, as light levels would be intermediate between open areas and underneath the
canopy. The process of natural regeneration is strongly influenced by edge environments
due to the effects of vegetation cover on microclimate conditions (Asbjornsen et al.
2004). Survival of Portulacaria afra, a keystone species in Thicket, is highest at the edge
of Thicket, due to protection from browsers and harsh climatic conditions in the open, as
well as less competition from established plants (De Ridder 2006). Even if species could
survive the harsh conditions of open areas (La Cock 2003), photosynthesis in Thicket
species is saturated at half of ambient light conditions (Holmes & Cowling 1993),
limiting the benefit that would be received from establishing in the open, as opposed to
the edge. Openings in mesic Thicket may allow the establishment of drought intolerant
species, although grass and shrub cover is much greater compared to xeric Thicket
(Everard 1987), greatly increasing competition.

The effectiveness of the edge as a regeneration niche may be diminished by climatic
events such as drought (Asbjornsen et al. 2004). The establishment of the woody
seedlings would be limited by soil moisture in Thicket, whereas the endemic succulent A.
africana would be less limited. Even though seedlings of shade intolerant species grow
faster than shade tolerant species, always in the open and possibly in shade, survival
underneath the canopy is lower (Kitajima 1994, Lin et al. 2002). For seedling
regeneration of other shade intolerant species to occur, resulting in their persistence in
101
Thicket, gaps as well as the edge of Thicket bushclumps need to be preserved (Holmes &
Cowling 1993).

The four species in this study would establish at different stages in the regeneration of
Thicket. Aloe africana, and to a less degree Ptaeroxylon obliquum, the most drought
tolerant species, could establish in the open and act as nurse species (La Cock 1992,
Watson 2002). The embryonic Thicket clump would then attract birds, by providing
cover as well as a perch (Vlok & Euston-Brown 2002). This will lead to the
establishment of Rhus dentata, a bird-dispersed species that can tolerate intermediate
environmental conditions. The bushclump will then be expanded, resulting in the further
amelioration of the undercanopy microclimate, until conditions favour the establishment
of Burchellia bubalina. All four species could then theoretically co-exist as they are able
to tolerate shade, with future establishment of the former species occurring at the edge of
Thicket clumps.

4.4.5 Conclusion
To effectively manage ecosystems for reestablishment of species, there is a need to
understand the complex and often synergistic interactions between different plant species,
the edge environment, and climatic variability that strongly influence regeneration
processes (Asbjornsen et al. 2004). This study focussed on the effect of the light and soil
water on the growth and survival of Thicket species.

Water supply is the dominant control of growth and seedling survival in Thicket species.
The three woody species that occur in Forest and Savanna outside of Thicket were able to
grow faster than the succulent and endemic Aloe africana, rejecting the first hypothesis.
However, this was a result of the generally low growth rates of drought tolerant species,
and not the superior response of drought intolerant species. All species in this study were
shade tolerant and able to grow in 80% shade, rejecting the second hypothesis that growth
would be highest in intermediate light conditions. One limitation is that shade could be
greater than 80% of ambient light, possibly resulting in reduced growth rates in Rhus
102
dentata and Ptaeroxylon obliquum. The observation that seedlings are limited to Thicket
clumps is a result of the established canopy facilitating the establishment of new
individuals, by protecting the soil underneath the canopy, and preserving the moisture
content of the soil. The low number of seedlings may be a result of a general dry climate
(Midgley & Cowling 1993), with seedling regeneration limited to Thicket types that
receive higher rainfall.

103
Chapter 5 Conclusion

Degradation and desertification occurs rapidly in semi-arid environments, and recovery is
generally slow as plant growth is limited by rain and denudation rainfall effectivity
(Wigand et al. 1995). In the Eastern Cape, large areas of Thicket have been degraded or
completely removed (Hoffman & Everard 1987, Palmer et al. 2006), resulting in a
transformed landscape that is less beneficial to nature, as well as to humans (Lechmere-
Oertel 2003). As natural regeneration has not been observed to any appreciable extent,
restoration efforts are necessary to return Thicket to a productive and valuable state
(Aucamp & Tainton 1984). For successful restoration, knowledge of seed biology is
essential to understanding community processes such as plant establishment, succession,
and natural regeneration (Vasquez-Yanes & Orozco-Segovia 1993). It is also critical for
the rehabilitation of ecosystems (Bell et al. 1993). The aim of this study was to further
the understanding of seed and seedling dynamics in Thicket, and in this way contribute to
its successful regeneration.

Plants are most vulnerable in their life histories during the seed germination and seedling
establishment phases (Vandvik & Vange 2003). This vulnerability originates from
seedlings being smaller, less competitive, less protected against predators, and more
susceptible to unfavourable environmental conditions compared to fully grown plants
(Vandvik & Vange 2003). Seedling survival may be greatly increased by avoiding times
and places where mortality risk is high (Fenner & Thompson 205). This may occur if
germination that is sensitive to environmental cues are selected for, as well as dispersal
mechanisms that will deposit seeds into safe sites (Gutterman 1994). The improved
ability to tolerate the dominant environmental stressors, such as drought and shade
resistance, will also promote establishment of seedlings. Thus the recovery of Thicket in
areas that have been transformed is most dependent on recruitment by the successful
germination of seeds and survival of seedlings in a relatively hostile environment. Where
Thicket has not been degraded or overutilised, it is maintained through vegetative
regeneration (Midgley 1991).
104
This study has contributed to the understanding of seed germination and seedling
dynamics of Thicket focussing on the higher rainfall Thicket types. The focus of this
study was the dominant germination and dormancy mechanisms of woody and succulent
Thicket species, the germinable seed bank, as well as the response of seedlings to drought
and shade. These topics were then related to seed dispersal, as well as the regeneration
niche of species.

All forms of Thicket (dune, mainland mesic and mainland xeric) lack a persistent
germinable seed bank (Pierce & Cowling 1991, Sigwela 2004, chapter 2). This severely
limits the rate of regeneration of vegetation (Van der Valk & Pederson 1989), as recovery
will only occur once seeds have been dispersed back into an area (Toh et al. 1999). As
the natural disturbance regime seldom results in the complete destruction of aboveground
biomass (Kerley et al. 1995), limiting the number of gaps that would not be quickly filled
by the vegetative expansion of surrounding plant individuals, a germinable seed bank
may not be a beneficial regeneration strategy in Thicket. This is supported by the absence
of difference between the seed bank of transformed and untransformed Thicket (Sigwela
2004).

As the seed bank cannot be relied upon for the regeneration of Thicket, the germination
requirements of recently dispersed seeds must be explored and understood. Hard seeded
species were found to respond to hot water to break seed coat-imposed dormancy, which
is equivalent in the environment to imbibition during periods of high temperatures in the
open. Few other species exhibited seed dormancy, and can thus be qualified as quiescent,
with only shade having a large effect on germination. Even though low light conditions
are usually a requirement for species that require burial for germination (Garwood 1989),
and the species that responded significantly to shade (Burchellia bubalina) had very
small seeds, low light conditions under the canopy (1% of ambient light) may be just as
effective. As most species do not have specific dormancy breaking requirements,
germination should not be a hindrance to the supply of seedlings. Germinability of seeds,
as well as seedling growth and survival, needs to be taken into account when identifying
105
possible nurse species, as well as identifying pioneer species in the succession of
Thicket (Vlok & Euston-Brown 2002).

Once it is established that germination should not be an impediment to regeneration, as
long as soil moisture is not limiting, knowledge of the tolerance of seedlings to prevailing
environmental conditions, as well as their survival in the natural environment, is
essential. Thicket species were found to respond differently to shade and drought,
indicating that species may have various regeneration niches, as some would be able to
establish on the edge of Thicket, as well as in openings in mesic Thicket, whereas other
species will only establish underneath the canopy. This has important implications for
rehabilitation, as well as developing an understanding of the assembly rules of Thicket
(Vlok & Euston-Brown 2002). Pioneer species, as well as species that benefit from less
shade (Holmes & Cowling 1993), would establish in openings in mesic Thicket, as well
as at the edge of existing Thicket clumps, whereas climax Thicket species would
establish in the centre of these clumps (Fabricius 1997). For the revegetation of degraded
sites, nurse plants capable of establishing under poor conditions, need to be dispersed
first into an area, and only then would other species be able to establish. The restoration
of the full diversity of Thicket species to a site would require all the supply of their
various regeneration niches.

Endozoochory could have a more important function of depositing the seeds in the most
beneficial environment, rather than promoting germination per se. Thus most plant
species may be adapted to endozoochory to improve seedling establishment and survival.
Passage through the gut of mammals seldom inhibits the germination of species, but
usually has a neutral or positive effect (Davis 2007, Traveset 1998). Dispersal by birds is
beneficial, as they forage preferentially in dense bush (Alcantara et al. 2000), resulting in
seeds being deposited in the ameliorated conditions underneath the canopy. This may not
be advantageous in vegetation such as tropical forest, where soil moisture is not limiting,
and light is severely reduced underneath the canopy, but in seasonal and semi-arid
environments, such as Thicket and Mediterranean shrublands (Alcantara et al. 2000),
establishment is preferential below the canopy. This is consistent with the finding that
106
many Thicket species, with the exception of Acacia karroo, a pioneer species (Hall
2003), lack specific dormancy mechanisms that promote establishment in canopy gaps or
open areas. Many Thicket species are shade tolerant, and are able to establish in the
microhabitat underneath the canopy.

This study supports the prediction that seedling regeneration is greater in Thicket that
receives higher rainfall, compared to xeric forms where vegetative regeneration
dominates (Sigwela 2004). However species richness and seedling density was low.
Species richness and density was greater in Buffels Thicket, where rainfall is above 900
mm, as opposed to other Thicket types that receive rainfall of less than 450 mm. The
proportion of woody seedlings was also much greater. Thus Thicket should be able to
regenerate naturally in moist climates. However, little to no regeneration of woody
species were observed in Sundays Thicket, as well as Shamwari (Albany Thicket 2) and
Loerie (Gamtoos Thicket 2). Both of these sites received lower rainfall than the others of
the same vegetation type, evidence that rainfall may be the dominant factor for seedling
survival.

Seed germination and seedling survival may be site specific and opportunistic,
responding to rainfall, similar to arid systems (Gutterman 1994). As rainfall is relatively
unreliable (Cowling 1984), using it as a dormancy breaking mechanism may not be
beneficial, especially if seeds do not remain viable for a protracted period of time.
Instead, a possible strategy whereby seeds germinate slowly and irregularly may allow
the plant to strike it lucky and geminate during a season of above-average rainfall, a
possible form of bet hedging (Baskin & Baskin 1998). Seeds with thicker seed coats
usually germinate slower, as observed in this study, but the seedlings of many species are
slow growing and have a large root biomass soon after germinating, similar to the tropical
dry forests of Ghana (Lieberman & Li 1992).

Even though regeneration may occur naturally in more mesic Thickets, a lack of dispersal
to safe sites may occur in mammal-dispersed species, as most indigenous mammals
have been replaced by livestock (Kerley et al. 1995), which are inferior in terms of seed
107
dispersal of Thicket species (Sigwela 1999). Although birds continue to be effective and
efficient dispersers (Sigwela 2004), some plant species have exhibited a specific disperser
requirement, resulting in poor dispersal in areas where livestock dominate. Therefore, a
lack of dispersers may explain the low number of seedlings in mesic Thicket, and poor
establishment conditions may be the cause of poor recruitment in xeric Thicket.
However, vertebrate-dispersed seeds were found in the seed bank of open areas.
Although mammals may spend much of their time in the dense cover, they do need to
move between Thicket clumps (Davis 2007), resulting in the dispersal of seeds into
openings. It was observed in this study that some gaps contained latrines that improve
establishment conditions for seedlings (La Cock 1992), thus facilitating their
establishment within small gaps formed by herbivores in Thicket (Holmes & Cowling
1993, Kerley et al. 1995).

Sigwela (2004) hypothesises that seedlings in Thicket remain in the seedling bank until
favourable conditions trigger their incorporation into the plant population. This was
contrary to what was observed in this study, as seedlings were found to be both shade and
drought tolerant (chapter 4), removing the mechanism to suppress seedling recruitment
into the canopy. Seedlings have a strong response to increased light (Marks & Gardescu
1998), a response that seems unlikely in the drought-resistant, slow growing species that
dominate the Thicket canopy. However, this is circumstantial and long-term field
measurements of seedling growth would be the only way of substantiating this
speculation.

All information can be integrated and utilised to prepare a rehabilitation and restoration
program in mesic Thicket. Many similarities exist between degraded subtropical
rainforest in Australia (Toh et al. 1999) and Thicket, and useful insights made by the
authors regarding regeneration and restoration can be compared and applied to Thicket.
Similar to Thicket, soil seed banks under isolated remnant trees in old fields were poor in
woody species, and most species are reliant on dispersal by birds or bats for recruitment
(Toh et al. 1999). Thus the availability of perches is an important factor in the rate of
recovery. The ability of various growth forms to act as perches and nurse species was
108
also investigated. Short, low growing trees are able to attract birds, but taller trees, as well
as a cluster of two or more trees, improves attractiveness to birds as perches. No
difference in seedling density was observed beneath the canopies of wind-dispersed and
animal-dispersed species (Toh et al. 1999). Most seedlings in Thicket were found under a
wind-dispersed species in the Fish River valley (La Cock 1992), whereas seedlings were
most common under a bird-dispersed species in Sundays Thicket (Sigwela 2004). No
seedlings were found in the open, few under low trees and shrubs, and most under tall
remnant species (Toh et al. 1999). A warning about assuming successful regeneration is
also given, as most seedlings identified were secondary, rather than primary forest
species, and very few survived to grow taller than 150 cm. A large number of species
were observed in the small seedling stage, but diversity decreased with size, especially in
late successional species. Very little recovery had occurred in 25 years, mainly due to a
lack of a persistent seed bank, as well as little dispersal as a result of a lack of suitable
perches for birds (Toh et al. 1999).

Guidelines proposed stipulate that restoration should begin with the early establishment
of poorly dispersed species, usually large-seeded ones, which should be planted in
clumps (Toh et al. 1999). Relating these restoration guidelines to Thicket, planting of
small clumps of fast-growing but drought-resistant species, that act as nurse plants to
seedlings (La Cock 1992, Sigwela 2004), as well as species preferred by local birds as
perches, is recommended. These species should have recurved branches (Vlok & Euston-
Brown 2002) to allow the formation of a microhabitat that is protected from herbivores
and able to accumulate litter, and maintain soil moisture. Seeds may be sown to
accelerate regeneration, although if active dispersal is occurring, this may be
unnecessary. Aspects that can be explored include the optimal distance between clumps,
as well as the effect of distance from intact Thicket on seed dispersal (Bruton 1992).
However, any restoration project would have to be long-term, as the dominant species in
semi-arid systems are long-lived, and changes in species composition may take longer
than human life spans (Wigand et al. 1995).

109
5.1 Opportunities for future research
Similar to many ecological studies, the main shortcoming of this study was the short
duration of the study. One of the recommendations of a seed bank study is sampling
during the different seasons, to identify whether seeds remain viable in the soil to form a
persistent seed bank, or whether they are only capable of forming transient seed banks. A
one year seed bank study with sampling occurring in the various seasons from the same
Thicket clumps would be ideal. However, this study must be done together with the
monitoring of which Thicket species are fruiting, as aseasonal fruiting is common
(Bruton 1992).

Research into the viability of Thicket seeds in the natural environment would further
contribute to the seed dynamics of Thicket. Seed burial experiments could be done to
determine how long seeds remain viable in the soil. Seeds can be buried in mesh bags,
both underneath the canopy as well as in the open, and periodically unearthed and
germinated.

The population dynamics of Thicket species has been neglected. To remedy this,
permanent quadrats could be set up and seedlings identified and tagged, and long term
monitoring can be done, to identify recruitment into the sapling and adult stages. Seeds
can be buried in the soil at the correct germination depth, and germination and seedling
survival can be monitored.

Thicket accumulates a lot of litter underneath the canopy. Litter may be beneficial or
detrimental to seedling establishment. The effects of litter can be investigated both in the
field, by comparing seedling richness and diversity in Thicket clumps that collect litter to
clumps devoid of litter, as well as in the glasshouse, where seed germination trials can be
done underneath varying depths of litter. The effect of litter from different species, both
succulent and woody growth forms, can be studied, especially for Portulacaria afra. An
excessive amount of litter is able to accumulate under the canopy of spekboom, and little
moisture seems to penetrate the canopy (pers. obs), possibly resulting in a hostile
microhabitat for seedlings.
110

As the risk of herbivory is greatest in the very early stages of establishment (Fenner &
Thompson 2005), the effect of defoliation through herbivory on seedling survival would
be another interesting topic of research. It was observed that some germinated seedlings
in the glasshouse were predated upon by insects or snails. While this resulted in the
mortality of some seedlings, others were able to grow new leaves, even though they were
still at the first leaf pair stage.

The main solution to Thickets problems may be the removal of man. However, as this is
not probable, and considering the damage already being done, the future of Thicket,
especially in the inland drier forms, is bleak without intervention, and active
rehabilitation and restoration is required urgently. The information gained in this study
will hopefully contribute to the maintenance and preservation of Thicket for many
generations to come.

111
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127
Appendix 1
Species found in the aboveground vegetation (V) and seedlings (S) of the study sites. BT = Buffels Thicket; AT = Albany Thicket;
ST = Sundays Thicket; GT = Gamtoos Thicket.
FAMILY SPECIES BT1 BT2 BT3 AT1 AT2 ST1 ST2 GT1 GT2
Acanthaceae Hypoestes aristata (Vahl) Sol. ex Roem. & Schult. V V
Acanthaceae Hypoestes forskaolii (Vahl) R.Br. V V V V V V V
Acanthaceae Hypoestes sp. S S S S S S
Aizoaceae Galenia pubescens(Eckl. & Zeyh.) Druce S
Anacardiaceae Rhus chirindensis Baker f. S S S
Anacardiaceae Rhus crenata Thunb. V
Anacardiaceae Rhus dentataThunb. V V
Anacardiaceae Rhus glauca Thunb. V V
Anacardiaceae Rhus laevigata L. V
Anacardiaceae Rhus longispina Eckl. & Zeyh. V V V
Anacardiaceae Rhus pallens Eckl. & Zeyh. V V V V V V
Anacardiaceae Rhus pterota C.Presl V V V V V V
Anacardiaceae Rhus sp. S S S s
Anacardiaceae Rhus sp2 V V V
Anacardiaceae Rhus tomentosa L. V V V V
Apocynaceae Acokanthera oppositifolia (Lam.) Codd V
Apocynaceae Carissa bispinosa (L.) Desf. ex Brenan V V V V V V V
Apocynaceae Cynanchum ellipticum (Harv.) R.A.Dyer V
Apocynaceae Fockea edulis (Thunb.) K.Schum. V V V V V
Apocynaceae Pachypodium bispinosum (L.f.) A.DC. V
Apocynaceae Sarcostemma viminale (L.) R.Br. subsp. viminale V S V V V V V S
Araliaceae Cussonia sp. S S S
Araliaceae Cussonia spicata Thunb. V V V V V
Araliaceae Cussonia thyrsiflora Thunb. V V
Asparagaceae Asparagus aethiopicus L. V V V
Asparagaceae Asparagus africanus Lam. V V V V V V V
128
Asparagaceae Asparagus crassicladus Jessop V V
Asparagaceae Asparagus racemosus Willd. V V V V V V
Asparagaceae Asparagus setaceus Kunth. V V
Asparagaceae Asparagus sp. S
Asparagaceae Asparagus striatus (L.f.) Thunb. V V V
Asparagaceae Asparagus suaveolens Burch. V V V V V
Asparagaceae Asparagus subulatus Thunb. V V V
Asparagaceae Asparagus volubilus Thunb. V V V
Asphodelaceae Aloe africana Mill. V S V V V V
Asphodelaceae Aloe ciliaris Haw. V V
Asphodelaceae Aloe ferox Mill. S V S
Asphodelaceae Aloe pluridens Haw. V S V S V V
Asphodelaceae Aloe striata Haw. V
Asphodelaceae Bulbine frutescens (L.) Willd. V
Asphodelaceae Gasteria bicolor Haw. var. bicolor V S V S S
Asteraceae Brachylaena discolor DC. V
Asteraceae Brachylaena elliptica (Thunb.) DC. V
Asteraceae Brachylaenia ilicifolia (Lam.) Phill. & Schweick V V V V
Asteraceae Chrysanthemoides monilifera (L.) T.Norl. subsp. monilifera V
Asteraceae Chrysocoma rigidula (DC.) Ehr.Bayer V
Asteraceae Gerbera piloselloides (L.) Cass. V
Asteraceae Helichrysum cymosum (L.) D.Don. subsp. cymosum V V S V S V
Asteraceae Pentzia incana (Thunb.) Kuntze V
Asteraceae Pseudognaphalium luteo-album (L.) Hilliard & B.R. Burtt S S
Asteraceae Pteronia incana (Burm.) DC. V V
Asteraceae Senecio angulatus L.f. V V V V
Asteraceae Senecio sp. V
Asteraceae Senecio inaequidens DC. V
Asteraceae Senecio pterophorus DC. S
Asteraceae Senecio radicans (L.f.) Sch.Bip. S V S V
Bignoniaceae Tecoma capensis Thunb.) Lindl. V S V V
Cactaceae *Opuntia aurantiaca Lindley V
129
Cactaceae *Opuntia ficus-indica (L.) Mill. V V V V V
Capparaceae Cadaba aphylla (Thunb.) Wild V
Capparaceae Capparis sepiaria L. var. citrifolia (Lam.) Tlken V V V V V V
Capparaceae Maerua cafra (DC.) Pax V V
Celastraceae Cassine peragua L. S S
Celastraceae Gymnosporia capitata (E.Mey. ex Sond.) Loes. V V V V V
Celastraceae Gymnosporia heterophylla (Eckl. & Zeyh.) Loes. V V V
Celastraceae Gymnosporia sp. S
Celastraceae Lauridia tetragona (L.f.) R.H.Archer V V V
Celastraceae Mystroxylon aethiopicum (Thunb.) Loes. subsp. aethiopicum S V V V V
Celastraceae Pterocelastrus tricuspidatus (Lam.) Walp. V V
Celastraceae Putterlickia pyracantha (L.) Szyszyl. V V S V V
Celtidaceae Celtis africana L. S
Chenopodiaceae Atriplex semibaccata R.Br. V
Commelinaceae Commelina africana L. V S V V V S V
Commelinaceae Commelina benghalensis L. V
Commelinaceae Cyanotis speciosa (L.f.) Hassk. V
Convolvulaceae Convolvulus sagittatus Thunb. V V
Crassulaceae Cotyledon velutina Hook.f. V V V V
Crassulaceae Crassula cordata Thunb. V V V
Crassulaceae Crassula cotyledonis Thunb. V V V
Crassulaceae Crassula expansa Dryand. S V S S
Crassulaceae Crassula mesembryanthoides (Haw.) D.Dietr. V V V
Crassulaceae Crassula ovata (Mill.) Druce V V V S V S
Crassulaceae Crassula perforata Thunb. V V V S
Crassulaceae Crassula spathulata Thunb. S
Crassulaceae Crassula tetragona L. V V V V
Crassulaceae Kalanchoe rotundifolia (Haw.) Haw. V S
Cucurbitaceae Kedostris nana (Lam.) Cogn. S V V V V S
Dracaenaceae Sansevieria aethiopica (L.) Druce V S V S V V V
Ebenaceae Diospyros dichrophylla (Gand.) De Winter V V S V S V
Ebenaceae Euclea sp. S S S
130
Ebenaceae Euclea undulata Thunb. V V V V V V
Euphorbiaceae Acalypha villicaulis Hochst. S
Euphorbiaceae Clutia daphnoides Lam. V V
Euphorbiaceae Euphorbia grandidens Haw. V V
Euphorbiaceae Euphorbia ledienii A.Burger V S V
Euphorbiaceae Euphorbia mammillaris L. V V
Euphorbiaceae Euphorbia mauritanica L. var. mauritanica V
Euphorbiaceae Euphorbia tetragona Haw. V
Euphorbiaceae Euphorbia triangularis Desf. V V S
Fabaceae Acacia karroo Hayne V V V S V V V
Fabaceae Schotia afra (L.) Thunb. var. afra V V S V V V V
Fabaceae Schotia latifolia Jacq.
Flacourtiaceae Dovyalis rhamnoides (Burch. Ex DC.) Burch. & Harv. S S v v v
Flacourtiaceae Scolopia mundii (Eckl. & Zeyh.) Warb. V V
Flacourtiaceae Trimeria grandifolia Hochst. Warb. V
Gentianaceae Sebaea macrophylla Gilg S
Geraniaceae Pelargonium peltatum (L.) L'Hrit S V V V S
Geraniaceae Pelargonium reniforme Curtis V
Hyacinthaceae Drimia sp. V
Hyacinthaceae Ledebouria revoluta (L.f.) Jessop V S
Hyacinthaceae Ornithogalum tenuifolium F. Delaroche. subsp. tenuifolium S
Hypoxidaceae Hypoxis stellipilis Ker Gawl. V V S
Iridaceae Watsonia sp. V
Lamiaceae Plectranthus madagascariensis (Pers.) Benth. var. madagascariensis S V S
Lobeliaceae Cyphia sylvatica Eckl. V
Malvaceae Abutilon sonneratianum (Cav.) Sweet V V V V S V V S
Mesembryanthemaceae Aptenia cordifolia (L.f.) Schwantes S
Mesembryanthemaceae Carpobrotus edulis (L.) L.Bolus V
Mesembryanthemaceae Delosperma calycinum L.Bolus V V V
Mesembryanthemaceae Drosanthemum hispidum (L.) Schwantes V V
Mesembryanthemaceae Mesem1 V V
Mesembryanthemaceae Aptenia cordifolia (L.f.) Schwantes V V
131
Mesembryanthemaceae Ruschia cymbifolia (Haw.) L.Bolus. V
Myrsinaceae Rapanea melanophloes (L.) Mez V
Oleaceae Jasminum angulare Vahl S S V
Oleaceae Olea europaea L. subsp. africana (Mill.) P.S.Green V V V V
Orchidaceae Bonatea speciosa (L.f.) Willd. V
Plumbaginaceae Plumbago auriculata Lam. V V V S V
Poaceae Panicum deustum Thunb. V V V V V V V
Poaceae Panicum maximum Jacq. V
Poaceae Pentaschistis curvifolia (Schrad.) Stapf V
Polygalaceae Polygala myrtifolia L. V
Portulacaceae Portulacaria afra Jacq. V V
Pteridaceae Cheilanthes viridis (Forssk.) Swartz var. viridis V V V V V V
Ranunculaceae Clematis brachiata Thunb. V
Rhamnaceae Scutia myrtina (Burm.f.) Kurz V V V V V V V
Rhamnaceae Ziziphus mucronata Willd. V S V S
Rubiaceae Burchellia bubalina (L.f.) Simms V V S V S
Rubiaceae Canthium sp. S
Rubiaceae Galopina circaeoides Thunb. S S
Rutaceae Clausena anisata (Willd.) Hook.f. Benth. var. anisata V V
Rutaceae Ptaeroxylon obliquum (Thunb.) Radlk. V V V V
Rutaceae Vepris lanceolata Comm. Ex A.Juss V S
Rutaceae Zanthoxylum capense (Thunb.) Harv. V V V
Salvadoraceae Azima tetracantha Lam. V V S V V V S V
Santalaceae Rhoiacarpos capensis (Harv.) A.DC. V V V V V
Sapindaceae Hippobromus pauciflorus (L.f.) Radlk. V S V V S V V V
Sapindaceae Pappea capensis Eckl. & Zeyh. V V V V
Sapotaceae Sideroxylon inerme L. subsp. inerme S V V V V V V
Solanaceae Ehretia rigida (Thunb.) Druce subsp. rigida V S V S V V V V
Solanaceae Lycium ferocissimum Miers V V V V
Solanaceae Solanum rigescens Jacq. V
Solanaceae Solanum sp. S S
Sterculiaceae Hermannia althaeoides Link V V V V
132
Tiliaceae Grewia occidentalis L. V S V S V S V V S
Tiliaceae Grewia robusta Burch. V V V
Unknown seedling1 S
Unknown seedling2 S
Unknown seedling3 S
Unknown seedling4 S
Unknown seedling5 S
Unknown seedling6 S
Unknown unknown sp1 V
Viscaceae Viscum crassulae Eckl. & Zeyh. V
Viscaceae Viscum obovatum Harv. V
Viscaceae Viscum rotundifolium L.f. V V V V V
Vitaceae Rhoicissus digitata (L.f.) Gilg & Brandt V S
Vitaceae Rhoicissus tridentata (L.f.) Wild & R.B.Drumm. subsp. tridentata V V V V V V V V V
Zygophyllaceae Zygophyllum morgsana L. V V

133
Appendix 2
Species identified from the germinable seed bank of Mainland Thicket.
FAMILY SPECIES
Aizoaceae Aizoon glinoides L.f.
Aizoaceae Aizoon rigidum L.f.
Anacardiaceae Rhus sp.
Anacardiaceae Rhus chirendensis Baker f.
Apiaceae Centella asiatica (L.) Urb.
Apiaceae Centella sp.
Apocynaceae Cynanchum obtusifolium L.f.
Apocynaceae Gomphocarpus fruticosus (L.) Aiton f. subsp. Fruticosus
Araliaceae Cussonia spicata Thunb.
Asparagaceae Asparagus sp.
Asphodelaceae Bulbine frutescens (L.) Willd.
Asteraceae Arctotheca calendula (L.) Levyns
Asteraceae Berkheya bipinnitafida (Harv.) Roessler subsp. bipinnatifida
Asteraceae Berkheya rigida (Thunb.) Bolus & Wolley-Dod ex Adamson & T.M.Salter
Asteraceae Cineraria saxifraga DC.
Asteraceae Conyza scabrida DC.
Asteraceae Cotula heterocarpa DC.
Asteraceae Dimorphotheca sp.
Asteraceae Gamochaeta pennsylvanica (Willd.) Cabrera
Asteraceae Helichrysum cymosum (L.) D.Don
Asteraceae Hypochaeris radicata L.
Asteraceae Nidorella auriculata DC.
Asteraceae Pseudognaphalium luteo-album(L.) Hilliard & B.L.Burtt
134
Asteraceae Senecio deltoideus Less.
Asteraceae Senecio glutinosus Thunb.
Asteraceae Senecio inaequidens DC.
Asteraceae Senecio polyanthemoides Sch.Bip.
Asteraceae Senecio pterophorus DC.
Asteraceae Sonchus oleraceus L.
Asteraceae Taraxacum officinale Weber.
Asteraceae Troglophyton capillaceum (Thunb.) Hillard & B.L.Burtt subsp. capillaceum
Brassicaceae Lepidium africanum (Burm.f.) DC.
Buddlejaceae Buddleja saligna Willd.
Cactaceae Opuntia ficus-indica (L.) Mill.
Caryophyllaceae Polycarpon tetraphyllum (L.) L.
Celastraceae Putterlickia sp.
Celtidaceae Celtis africana Burm.f.
Chenopodiaceae Atriplex semibaccata R.Br.
Chenopodiaceae Atriplex vestita (Thunb.) Aellen
Chenopodiaceae Chenopodium album L.
Chenopodiaceae Chenopodium ambrosioides L.
Chenopodiaceae Chenopodium carinatum R.Br.
Chenopodiaceae Chenopodium murale L. var. murale
Commelinaceae Commelina africana L.
Commelinaceae Commelina benghalensis L.
Crassulaceae Crassula expansa Dryand.
Crassulaceae Crassula spathulata Thunb.
Crassulaceae Crassula tetragona L.
Crassulaceae Kalanchoe rotundifolia (Haw.) Haw.
Cyperaceae Ficinia sp.
Cyperaceae Kyllinga alata Rottb.
135
Cyperaceae Mariscus congestus (Vahl) C.B.Clarke
Ebenaceae Diospyros dicrophylla (Gand.) De Winter
Euphorbiaceae Acalypha villicaulis Hochst.
Euphorbiaceae Adenocline acuta (Thunb.) Baill.
Euphorbiaceae Euphorbia inaequilatera Sond. var. inaequilatera
Euphorbiaceae Phyllanthus incurvus Thunb.
Euphorbiaceae Phyllanthus verrucosus Thunb.
Fabaceae Acacia karroo Hayne
Fabaceae Acacia saligna (Labill.) H.L.Wendl.
Fabaceae Medicago polymorpha L.
Fabaceae Senna sp.
Gentianaceae Sebea macrophylla Gilg
Hyacinthaceae Ornithogalum tenuifolium F.Delaroche subsp. tenuifolium
Lamiaceae Plectranthus madagascariensis (Pers.) Benth.
Lamiaceae Salvia triangularis Thunb.
Lobeliaceae Cyphia sylvatica Eckl.
Lythraceae Lythraceae sp.
Malvaceae Abutilon sonneratianum (Cav.) Sweet
Malvaceae Hibiscus pusilla Sm.
Malvaceae Sida cordifolia L.
Malvaceae Sida ternata L.f.
Mesembryanthemaceae Aptenia cordifolia (L.f.) Schwantes
Mesembryanthemaceae Carpobrotus sp.
Mesembryanthemaceae Delosperma ecklonis (Salm-Dyck) Schwantes
Mesembryanthemaceae Drosanthemum sp.
Mesembryanthemaceae Mesembryanthemum aitonis Jacq.
Molluginaceae Pharnaceum sp.
Oxalidaceae Oxalis caprina L.
136
Oxalidaceae Oxalis incarnata L.
Oxalidaceae Oxalis punctata L.f.
Oxalidaceae Oxalis smithiana Eckl. & Zeyh.
Poaceae Cynodon dactylon (L.) Pers.
Poaceae Digitaria natalensis Stent
Poaceae Digitaria sanguinalis (L.) Scop.
Poaceae Ehrharta erecta Lam.
Poaceae Eustachys paspaloides (Vahl) Lanza & Mattei
Poaceae Panicum deustum Thunb.
Poaceae Panicum maximum Jacq.
Poaceae Paspalum dilatatum Poir.
Poaceae Paspalum distichum L.
Poaceae Poa annua L.
Poaceae Sporobolus africanus (Poir.) Robyns & Tournay
Poaceae Sporobolus fimbriatus (Trin.) Ness
Ranunculaceae Clematis brachiata Thunb.
Rubiaceae Anthospermum sp.
Rubiaceae Galopina circaeoides Thunb.
Rubiaceae Richardia brasiliensis Gomes
Rubiaceae Spermacoce natalensis Hochst.
Rutaceae Vepris lanceolata (Lam.) G.Don
Sapindaceae Sideroxylon inerme L. subsp. inerme
Scrophulariaceae Nemesia affinis Benth.
Scrophulariaceae Phyllopodium diffusum Benth.
Scrophulariaceae Selago corymbosa L.
Scrophulariaceae Sutera campanulata (Benth.) Kuntze
Scrophulariaceae Sutera polyantha (Benth.) Kuntze
Solanaceae Lycium ferocissimum Miers
137
Solanaceae Nicotiana glauca Graham
Solanaceae Solanum lichtensteinii Willd.
Solanaceae Solanum nigrum L.
Solanaceae Solanum retroflexum Dunal
Tiliaceae Grewia occidentalis L. var. occidentalis
Verbenaceae Lantana camara L.
Verbenaceae Priva meyeri Jaub. & Spach var. meyeri

138
Appendix 3
Mean (n = 4) Frequencies, Growth form (G) and dispersal mechanism (D) of individuals per plant species (m
-2
) in the germinable seed bank underneath
the canopy (C), at the edge (E) and in openings (O) of all sites. See text for site codes. (Wo = woody, He = herb, S = succulent, Gr = graminoid, Cl =
climber, Ge = geophyte. A = autochory, W = anemochory, Z = zoochory)
SPECIES G D MKT1 MKT2 MKT3 XKT1 XKT2 XKT3 MST1 MST2 MST3
C E O C E O C E O C E O C E O C E O C E O C E O C E O
Abutilon sonneratianum Wo W 16 8 32 8
Acacia karroo Wo Z 16 8 16 8 8
Acacia saligna Wo Z 8
Acalypha villicaulis He A 4 8 16 16 48 24 8
Adenocline acuta He A 8
Aizoon glinoides S A 8 8 8 8 8
Aizoon rigidum S A 24
Anthospermum sp. Wo Z 8
Aptenia cordifolia S Z

48 16
Arctotheca calendula He W 8 8 8 8
Asparagus sp. Cl Z 8
Atriplex semibaccata Wo Z 8
Atriplex vestita Wo W 8
Berkheya bipinnitafida
subsp bipinnatifida
He W 8 8 8
Berkheya rigida He W 32 64 16 8
Buddleja saligna Wo
A,
Z
32 16
Bulbine frutescens S
A,
W
8
Carpobrotus S Z 16
Celtis africana Wo Z 16 8
Centella asiatica He Z 8 24 16 14 16 16 72
139
Centella sp. He Z 24 24
Chenopodium album He W 24 16
Chenopodium
ambrosioides
He W 8 72 8 8
Chenopodium carinatum He W 8
Chenopodium murale He W
Cineraria saxifraga Wo W 8 16 4
Clematis brachiata Cl W 8
Commelina africana He Z 8
Commelina
benghalensis
He Z 8
Conyza scabrida Wo W
Cotula heterocarpa He W 8 8 8
Crassula expansa S W 48 24 28
42
4
4 48 16 8 8 48 32
Crassula spathulata S W 8
Crassula tetragona S W 8 8 8
Cussonia spicata Wo Z 16
Cynanchum obtusifolium Cl W 8
Cynodon dactylon Gr W 16 24 4 24 8 8 48 16 8 24 8 12 64 64 72 16 32 24 32 8
Cyphia sylvatica Cl W 14 288
14
4
12
8
296 48
Delosperma ecklonis S A 8 8 8 8 8
Digitaria natalensis Gr W 8 24 72 24
Digitaria sanguinalis Gr W 8 16 8 24 24
11
2
352 424 16
Dimorphotheca sp. He W 8
Diospyros dicrophylla Wo Z 8
Drosanthemum sp. S A 8
140
Ehrharta erecta Gr W 8 192 24 28 24 96
Euphorbia inaequilatera He A 48 344
23
2

37
6
728 34
17
6
48
111
2
936 92 616 8 32 4
Eustachys paspaloides Gr W 24
Ficinia sp. Gr W,Z 2 16 32 32 24 4
Galopina circaeoides He A 8 16 64 184
Gamochaeta
pennsylvanica
He W,Z 8 14 32 8 8 8 16 16 8 48 8 8 16
Gomphocarpus
fruticosus
He W 8 16
Grewia occidentalis Wo Z 8 8 8
Helichrysum cymosum Wo W,Z 8 24 4 48 64 8
Hibiscus pusillus He W
Hypochaeris radicata He W,Z 16
Kalanchoe rotundifolia S
A,
W
16 24
Kyllingia alata Gr W,Z 64
Lantana camara Wo Z 8
Lepidium africanum He W 8 32 8
Lycium ferocissimum Wo Z 8
Lythraceae sp. He W 4 16
Mariscus congestus Gr W,Z 8 8 32 64 24 24 12
Medicago polymorpha He Z 8 4 56
Mesembryanthemum
aitonis
S A 8 8 16 24 8
Nemesia affinis He W 8
16
8
16 8
41
6
38
4
28
Nicotiana glauca Wo
A,
W
16 24
Nidorella auriculata He W
Opuntia ficus-indica S Z 8
141
Ornithogalum
tenuifolium
subsp. tenuifolium
Ge A 8
Oxalis caprina Ge A 16
Oxalis incarnata Ge A 8
Oxalis punctata Ge A 8 8
Oxalis smithiana
Ge
o
A 8 8
Panicum deustum Gr W,Z 8 32 24 8 16 72 32 8 56 8 4
Panicum maximum Gr W,Z 24 4 16 16 24 8 8 184 56 12 72
18
4
21
6
96 56 32 14 12
16
8
64 96 16
Paspalum dilatatum Gr W 8
Paspalum distichum Gr W 32 72
Pharnaceum sp. He A 8
Phyllanthus incurvus He A 16 8 16 8 8
Phyllanthus verrucosus Wo A 24
Phyllopodium diffusum He W 8 16
Plectranthus
madagascariensis
He Z 8
Poa annua Gr W 8
Polycarpa tetraphyllum He W 16 8 168
Priva meyeri var meyeri He A,Z
19
2
376
Pseudognaphalium
luteo-album
He W,Z 48 192 96 8 24 8
Putterlickia sp. Wo A,Z 8
Rhus sp. Wo Z 8 8 8
Rhus chirendensis Wo Z 8
Richardia brasiliensis He A 8 8
Salvia triangularis He Z 8 8
Sebea macrophylla He A 8 8
142
Selago corymbosa Wo W 16 72 8 16 8
Senecio deltoideus Cl W 8
Senecio glutinosus He W 8 8 8 8
Senecio inaequidens He W 56 64 8 56 88 16 16 8 8
Senecio
polyanthemoides
He W 8 4 56 16 8 24 16 8 8
Senecio pterophorus He W 8 16 48 14
Senna sp. Wo Z 8 8
Sida cordifolia Wo W 8 8
Sida ternata He W 8 8
Sideroxylon inerme Wo Z 32
Solanum lichtensteinii Wo Z 8 8 8
Solanum nigrum He Z 8
Solanum retroflexum He Z 24 8 8
Sonchus oleraceus He W,Z 8 8 8 32 8 8 8 24
Spermacoce natalensis He A 96 136 72
Sporobolus africanus Gr A 72 136
26
4
8 56 4 8 8 8 8 24
Sporobolus fimbriatus Gr A 8
Sutera campanulata He W 8 8 8
Sutera polyantha He A 8 16 8 48 12 8
Taraxacum officinale He W,Z 8 24
Troglophyton
capillaceum
subsp. capillaceum
He
W,
Z
8
Vepris lanceolata Wo Z 8
Total
50
0
141
8
80
0
53
6
125
2
55
0
63
6
156
8
111
6
32
8
18
0
157
6
35
2
20
4
55
2
40
4
49
6
20
4
104
4
44
4
110
4
15
0
15
6
30
8
62
4
58
0
16
8

Weatherall ThomasCR

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Seed Dynamics and Seedling Survival in Mainland

Thicket of the Eastern Cape

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