Beruflich Dokumente
Kultur Dokumente
Faculty of Science
Department of Biological sciences
November 2008
Supervisor: Dr J.K. Mfune
Table of contents
Acknowledgement................................................................................................................. 2
Abstract................................................................................................................................. 3
1. Introduction ....................................................................................................................... 4
2. Literature review ............................................................................................................... 7
2.2 Descriptions of the Datura strammonium and Datura innoxia ..................................... 8
2.2.1 D. strammonium ....................................................................................................... 8
2.2.2 D. innoxia ................................................................................................................. 8
2.3 Factors that influence or encourage the invasion of the alien invasive species ............ 8
2.4 Problems or concerns of the Datura alien invasive species ......................................... 9
2.5 Preventative measures and eradication of alien invasive species................................ 9
3. Objectives ....................................................................................................................... 10
4. Key questions ................................................................................................................. 10
5. Hypothesis ...................................................................................................................... 10
6. Methodology ................................................................................................................... 11
6.1 Study sites ................................................................................................................ 11
6.2 Data collection and records ....................................................................................... 11
6.3 Data analysis............................................................................................................. 11
7. Results ............................................................................................................................ 12
8. Discussion ...................................................................................................................... 16
Site 1............................................................................................................................... 16
Site two ........................................................................................................................... 17
Relationship between crown width and reproductive structures....................................... 18
Ecological importance of different flowering patterns ....................................................... 18
9. Conclusions .................................................................................................................... 19
10. References.................................................................................................................... 20
11. Appendix ....................................................................................................................... 21
Acknowledgement
Firstly, thanks to the Almighty God for all his guidance and protection during the field work
up to the point of completing the report. Secondly, I would like to thank Dr J.K. Mfune my
supervisor for all the sacrifices and for always being so helpful in all times of need.
Thirdly, a warm thank you to UNAM students Atshikoto Eliaser and Haihambo Isai, for their
accompany and assistance during the field work. Last but not least, thanks to the statistic
department and all colleagues. Finally, more thanks to my family and friends for the words of
courage and motivation during the course of my project. May you all be blessed today and
forever! Thank you so very much
Abstract
D. innoxia and D. strammonium are some of the invasive alien species found in Namibia and belong
to family Solanaceae. Alien species refer to species which are introduced into a new geographical
area where it did not evolve. These species become invasive if they start spreading in ways that are
destructive to human interest and other native fauna and flora. They have different survival strategies
such as the ability to produce vast number and long-lived seeds, fast germination, rapid maturation,
growth and reproduction. The main objective of the study was to determine and compare the density
of reproductive structures of D. innoxia and D. strammonium, and determine their flowering patterns.
Two study sites along the Gammams river in Hochland Park in the Windhoek city were selected
based on the presence of D. innoxia and D. strammonium. Two quadrats, each 10m x 20m were laid
down, and data were collected during three occasions at the interval of two weeks between AprilJune 2008. Total number of buds, flowers, capsules (green and brown intact, green and brown
exploded) on each D. innoxia and D. strammonium, in each quadrat were recorded.
For normality test, Kolmogorov-Smirrnov test was used for buds, flowers, green intact capsules,
green exploded capsules, brown intact and brown exploded capsules (all t=0.00, p<0.05 values) were
obtained. T-test revealed that there was a significant difference between the following at site 1: buds
(t=7.7, df= 264, P < 0.05), flowers (t=8.66, df= 264, P < 0.05), green intact capsules (t=-4.18 df= 264,
P < 0.05), green exploded capsules (t=3.02 df= 264, P < 0.05) and brown exploded capsules (t= df=
264, P < 0.05 4.53). Except for brown intact pods where t=-1.37 P > 0.05, and df=264. This area was
0
located across the tarred road at 22 3415.4S, and 017 0413.7E and Alt: 1663 m ab.s.l) and water
0
was blocked from flowing across. At study site 2 (22 3425.8S and 017 0428.1E and Alt: 1666 m
ab.s.l) the t-test revealed that there was no significant difference between the buds (t = 0.99 df = 338
and P < 0.05.), flowers, (t = 0.23, df = 338 and P < 0.05), green intact capsules (t=0.77 df = 338 and
P < 0.05.), green exploded capsules (t = 2.65 df = 338 and P = 0.05.) and brown intact capsules (t = 1
df = 338 and P < 0.05.). Except only green exploded capsules where t = 3.09, df = 338 and P < 0.05.
There was no correlation between the crown width and the following: buds (r =0.321), flowers (r =
0.239454), green exploded capsules (r = 0.07644656), brown intact capsules (r = 0.178087), and
brown exploded capsules (r = 0.39382), except green intact pods (r = 0.567661) for D. innoxia. For D.
strammonium buds (r = 0.322336995), flowers (r = 0.278430477), green intact capsules (r =
0.267823), green exploded capsules (r = 0.09746044), brown intact capsules (r = 0.349538) and
brown exploded capsules (r = 0.373589).
The study concluded that two species have different densities of reproductive structures produced at
different time intervals which may probably depend on the physical structure and characteristics of
the habitat. On the basis of producing both reproductive structures at the same time and at different
time interval and still contained green leaves and branches, it was concluded that D. innoxia has a
continuous flowering pattern which was not the case with D. strammonium which reproduce, explode
once and dies, while D. innoxia is still flowering.
1. Introduction
Every species has its natural geographical range, some are cosmopolitan and some may be
endemic to specific locations (Brown and Gibson, 1983). Krebs (1994) indicated that,
transplant experiments are carried out to analyze whether the absence of some species at
particular areas is due to poor dispersal ability or other factors such as the failure of the
species to recognize the area as suitable or whether existence of certain efficient predators
or parasites in that area are responsible. During the transplant experiments, individuals are
moved from their native habitats to other areas, where they can be determined if they can
establish there or not.
Humans are responsible for introduction of various species from one areas of their natural
geographic range to new habitats for various reasons including agricultural, scientific, as
well as purely aesthetic beauty or for ornamental purposes. Henderson and Wells (2001)
defined alien species as the species with the ability and potential to establish in new
environments outside its natural distributions, were it occurs as a foreigner species. Alien
invasive are therefore species that human move intentionally or accidentally from their
native locations on to new geographical region and spread in ways that are destructive to
human interest and other native floral and fauna (Campell and Reece, 2005). But, which
areas are more prone to exotic invasions? Bullock et al., (2001) indicated that most studies
have concluded that the cultivated land, land much modified by human practices, protected
national parks, urban industrial areas, and habitats suffering from periodic disturbances are
more at risk. For example, Henderson and Wells (2001) stated that D. innoxia and D.
strammonium are mainly found at the edges of dams, invade riverbeds and mostly disturbed
areas.
Invasive alien species have affected native biota in almost every ecosystem type on earth
because of some unique characteristics they posses, and this became one of the worldwide
concerns. For instance, Bromilow (2001) indicated that D. strammonium produce poisonous
seedlings and human fatalities have been in cases where people have eaten it deliberately
or accidentally. There are three Datura species recorded in Namibia Datura innoxia (the
downy thorny apple), Datura strammonium (Common thorn apple) and Datura ferox. D.
innoxia is the most invasive although D. ferox and D. strammonium are also widespread.
These alien invasive plants belong to the same Solanaceae family and genus Datura, and
little information are known about them (Macdonald et al. 1986). High infestation of D.
innoxia was recorded from Avis dam, section of the Kuiseb river and rivers in the Hoanib
river flood plain (Macdonald et al. 1986). They originated from the south and Central
America and were introduced and dispersed unintentionally in different regions, including
Namibia (Cowling, Pierce and Richard, 1997).
About 40 species of alien invasive plants have been identified in Namibia such as Nicotiana
glauca, Opuntia species, Lantaria camara, Eucalyptus species and Datura species to
mention a few. Datura species show a world wide distribution, and their infestations are
widespread over the entire Namibia. Datura species are mostly prone to river washes areas
associated with perennial river systems e.g. the Okavango and Caprivi areas as shown by
Macdonald et al (1985). All of the Datura species are non-woody herbaceous plants, with
leafy green and bright pink to white flowers. Their seeds are found in small fruit which is
completely covered with short sharp spines. Their stalks are bristle, leaves are flat and can
either be multi - edged or ovoid (Thomson, 1978). D. innoxia and D. strammonium are
cultivated to obtain leaves and seeds. These plants contain poisonous substances, yet are
used in cigarettes and as smoke candles for asthmatics.
Brown et al. (1985) indicated that the greatest threat of alien invasive species in a given
area is posed by species, which originate from similar habitats in other parts of the world.
Generally, the smaller the number of individuals of alien species introduced into a country,
the smaller are its chances of becoming established as an invasive problem. Therefore, all
efforts at reducing the number of alien species should be regarded as worthwhile even if
total eradication of the species seems impossible Brown et al. (1985).
However, alien invasive species have general special characteristics which make them to
have a world wide distribution. They possess early sexual maturity due to rapid growth and
maturity, high reproductive output such as the production of many pods with vast seeds
even under harsh climatic conditions and other unfavorable conditions (Henderson and
Wells, 1987). Long life of their seeds, high ability to retrieve nutrients and water (H 2O), high
photosynthetic rate, lack of natural predators, pathogens and parasites contributes as well,
thus out-competing the indigenous competitors (Henderson and Wells, 1987). This may
eradicate the indigenous species such as the vegetations in their surrounding which may
change the ecosystem structure. Predators to the eradicated species can migrate to other
areas with no invasions, and humans may also be affected if the floral and fauna species in
their surrounding happens to be disturbed.
Recent studies have shown that Datura species begin reproducing at a very early vegetative
state, independent of the height or width. They begin to produce a bud, which develops into
the flower, beginning at the branch node which later develops into the spine fruit (Jansen,
2001). Some of the Datura species are self fertile (fertilization takes place using its own
pollen) and some are cross pollinated (fertilization of the flower by a pollen from a different
plant of the same species) (Jansen, 2001). Brown et al. (1985) stated that very little is
known about the reproductive strategies of the Datura species. It is important to know more
about the reproductive mechanism of the species, if one is to plan for the proper and
effective eradication strategies.
Greater understanding of their flowering patterns may allow one to determine if it is possible
for species that belong to the same genus and family to have similar or different flowering
strategies. This will help the conservationists, agriculturalists and even the systematists
formulate and initiate possible effective means that will lead to their eradication and or
prevent their manifestation. Again, one may determine not to eradicate it, provided they are
of some economic importance. One may desire to know conditions under which its negative
effects at a given site can be minimized. It is necessary for one to fully understand the
reproductive strategies of the invasive alien species for conservation of biotic diversity which
is one of the main aims of nature conservation authorities (Boyer and Boyer 1988).
The study by Katjirua (2008) indicated that D. innoxia was widely distributed in riverbanks,
disturbed areas and around human settlements in Namibia. Katjirua also indicated that
Datura innoxia is a perennial bushy herbaceous plant with a floral development but
mentioned nothing about the D. strammonium as the study was more on the distribution and
abundance of D. innoxia specifically. Therefore, the project mainly focused on the
reproductive strategies of the invasive alien D. innoxia and D. strammonium, but specifically
on the flowering patterns.
2. Literature review
Human migration has been occurring on large scales for several tens of thousands of years
and they have promoted processes of species transfer within continents, between
continents and nearby islands and between the continents themselves (Brown and Gibson
1983). Leveque and Mounolovu (2003) added that the voluntary or accidental migration of
individuals from one ecosystem to another is relatively a frequent phenomenon. Alien
invasive species can be determined by their biological and attributes to the dispersal
capabilities with the indigenous flora and fauna as shown by Venter (2002).
2.1 Types of Datura species and other alien invasive species in Namibia
Avery et al (1950) indicated that there are about eleven known Datura species such as D.
ferox, D. querlifolic, D. pruinosa, D. leichhardtii, D. meteloides, D. discolor, D. metel, D.
ceratocaule, D. wrightii, D. innoxia and D. strammonium. D. innoxia and D. strammonium
are widely found in Namibia. However, it was again recently found that there are actually
more than 11 Datura species described. These include the D. brugmansia, D. arbovea, D.
aurea, D. candida, D. dolichocarpa, D. fatuosa, D. indica, D. sanguinea, D. suaveolens, D.
vulcanicola, and D. willemsi in addition to the above.
Apart from the Datura innoxia and Datura strammonium, other examples of alien invasive
found in Namibia include the Nicotiana glauca, Opuntia species, Salvinia molest, Prosopis
species, Eucalyptus species. Opuntia ficus indica is the most important Opuntia species
which was introduced as a fodder plant for its edible fruit. Prosopis species were introduced
in Namibia in 1912 as fodder as well as shade trees. High infestation of Prosopis species
occur on municipal land near Windhoek, near Rehoboth, in Swakop river in the NamibNaukluft Park and in the vicinity of the Orange river. Moreover, Nicotiana glauca is one of
the most widespread species in the drier western regions of Namibia and along the length of
the Orange river. It has entered Namibia in horse fodder imported from South America
(Macdonald et al. 1986). Apart from D. innoxia and D. strammonium which are used to
obtain poisonous substances, other types of Datura species have several chemicals which
can be used as drugs in hospitals such as astropine (Thomson, 1978). Astropine is used by
opticians to dilate pupils and by doctors to treat heart and urinary problems. Some herbalists
use it to treat birth and menstrual pains
production. This will eventually reduce the number of alien species establish in a particular
area Macdonald and Jarman (1985).
3. Objectives
1. To determine and compare the density of buds, flowers and pods of D. innoxia and D.
strammonium growing in the selected areas of the city of Windhoek.
2. To determine and compare the flowering patterns of D. innoxia and D. strammonium in
selected areas in the city of Windhoek.
4. Key questions
1. Is there a difference in density of buds, flowers, and pods (green intact, green exploded,
brown intact and brown exploded) of D. innoxia and D. strammonium in selected areas of
Windhoek?
2. Is there a difference in the flowering patterns between the D. innoxia and D. strammonium
species in selected areas of the city of Windhoek?
5. Hypothesis
1. Ho: There is no significant difference in the density of buds, flowers and pods between the
invasive alien D. innoxia and D. strammonium in the selected areas of the city of Windhoek.
H1: There is a significant difference in the density of buds, flowers and pods between the
alien invasive D. innoxia and D. strammonium in the selected areas of the city of Windhoek.
2. Ho: There is no significant difference in the flowering patterns between the alien invasive
D. innoxia and D. strammonium in the selected areas of the city of Windhoek.
H1: There is a significant difference in the flowering patterns between the alien invasive D.
innoxia and the D. strammonium in the selected areas of the city of Windhoek.
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6. Methodology
6.1 Study sites
Two different sites were chosen for the study in Hochland Park and named as site 1 and site
2, and are along the Gammams River. Site 1 was opposite the tarred road (2203415.4S,
01700413.7E and Alt: = 1663 m ab.s.l), located at a rocky area which was dry. Site 2 was
situated next to the river which was still containing water (at 2203425.8 S and
01700428.1E at the altitude of 1666 m above sea level) and habitat soil was gray and
limited rocks were available. These sites were chosen because of the presence of both D.
innoxia and D. strammonium growing at the same areas (sites) together. Two quadrats of 20
x 10 m each were laid down along side the river bed at each site.
6.2 Data collection and records
In each quadrat, at each site the following were counted and recorded about the D. innoxia
and D. strammonium. Total number of buds, flowers, capsules (green intact, green
exploded, brown intact, and brown exploded). Selections of quadrats were based on the
presence of D. innoxia and D. strammonium on each site. These data in each quadrat were
recorded three times at an interval of two weeks. Each of the three recording sessions was
done within two days. Sampling was carried out between April and June 2008.
6.3 Data analysis
The Kolmogorov-Smirrnov test was used to test whether all the data (of buds, flowers, green
intact capsules, green exploded capsules, brown intact and brown intact capsules) collected
for D. innoxia and D. strammonium were normally distributed. The descriptive statistics were
done to obtain the summary statistics of the mean density of all buds, flowers, capsules
(green intact, green exploded, brown intact and brown exploded) on each D. innoxia and D.
strammonium in each quadrat and at each habitat. The independent two sample t-test was
used to determine whether there was a significant difference in the mean density of the
buds, flowers, and capsules (green intact, green exploded ,brown intact and brown
exploded) of D. innoxia and the D. strammonium in the selected areas of Windhoek.
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7. Results
The mean density of the reproductive structures of D. innoxia and D. strammonium during
the three sampling times in Hochland park site 1 are represented in figure 1 below.
Figure 1: Mean density (no/plant) of the reproductive structures of the alien invasive D.
innoxia and D. strammonium species in Hochland park (site 1) all three sampling times. The
reproductive structures are represented by the mean number of buds, flowers, green intact
12
and green exploded capsules, brown intact and brown exploded capsules respectively. Bar
on each histogram indicates the standard error of the mean.
Figure.1. A represents first sampling time, Figure 1.B, second sampling time after two weeks
and Figure 1.C, third sampling time after two weeks. D. innoxia has a high density of buds
and flowers but D. strammonium has a high density of green intact, green exploded and
brown exploded capsules during all the three sampling times. There was no any bud and
flower on D. strammonium and has a very low density of green intact and brown intact pods
compared to a high density of brown exploded pods at the same sampling time (C).
There was a significant difference between most of the reproductive structures. Buds (t=7.7
P < 0 05 and df = 264), flowers (t = 8.66 P < 0 05 and df = 264), green intact pods (t = 4.18
P < 0 05 and df = 264), green exploded pods (t=3.02 P < 0 05 and df = 264) and brown
exploded pods (t= 4.53 P < 0 05 and df = 264) where P < 0 05 and df = 264 for all. Except
for brown intact pods where t=-1.37 P > 0.05, and df = 264.
13
The mean density of the reproductive structures of D. innoxia and D. strammonium during
the three sampling times at site 2 are represented in Figures 2 below.
Figure 2: Mean density (no/plant) of the reproductive structures of the alien invasive D.
innoxia and D. strammonium species in Hochland Park (site 2) for three sampling times. The
reproductive structures are represented by the mean number of buds, flowers, green intact
and green exploded capsules, brown intact and brown exploded capsules respectively. Bar
on each histogram indicates the standard error of the mean.
Figure 2. A represents first sampling time, Figure 2 B second sampling time after two weeks
and Figure 2. C third sampling time after two weeks. D. strammonium has a high density of
14
flowers and green intact capsules compared to D. innoxia, and none of the species has
started producing green intact, brown intact and brown exploded capsules both during the
first and second sampling time. In Figure 2 C, D. strammonium started producing green
exploded and brown exploded capsules.
There was no significant difference between buds (t=0.99, 338 P> 0.05), flowers (t=0.23 df =
338 and P < 0.05), green intact capsules (t=0.77 df = 338 and P < 0.05), green exploded
capsules (t=2.65 df = 338 and P < 0.05) and brown intact capsules (t=1 df = 338 and P <
0.05), where P > 0.05 and df=338. Except only green exploded pods where t= 3.09, df =
338 and P < 0.05. From the graph, there was a high density of buds, flowers up to the brown
exploded capsules on D. strammonium. Extremely low density of green exploded capsules
on D. innoxia and none of the brown intact and brown exploded.
Table1. The correlation coefficient (r) between the reproductive structures and crown width
of alien invasive D. innoxia and D. strammonium in Hochland park
Reproductive
D. strammonium
Buds
0.321240
0.32233699
Flowers
0.239454
0.278430477
0.567991
0.267823
0.07644655
0.097462044
0.178087
0.349538
0.39382
0.373589
There was no correlation between the reproductive structures and the crown width of D.
innoxia and D. strammonium. Except only for the green intact capsules on D. innoxia.
15
8. Discussion
Site 1
Both the D. innoxia and D. strammonium in site 1 have buds, flowers and green intact pods,
where D. innoxia has high density of the buds and flowers as shown in figure 1. This
indicates that the two species do flower at the same time. It can also happen as this might
be the right season where the two species flower, as April is also a rain season in Namibia,
thus providing water and increasing seed germination and plant growth. High density of
green intact, green exploded capsules, brown exploded capsules on D. strammonium as
indicated in figure 1 for site one and two gives an indication that most of the flowers on D.
strammonium grow and mature faster than that of the D. innoxia.
This allows the green intact capsules to mature and develop to green exploded and brown
exploded capsules, and then the plant dies, while the D. innoxia is still alive and has buds
and flowers at the same time. The faster maturity of D. strammonium can be due to the fact
that it may have a high rate of nutrient and water uptake than the D. innoxia which then lead
to the faster growth and early maturity to produce buds, flowers, green intact capsules,
green exploded capsules, brown intact and brown exploded capsule respectively before
drying up as mentioned in literature (Henderson and Wells, 1987). That is why the density of
buds, flowers, green intact and green exploded capsules on D.strammonium at habitat one
decreased so much during the third sampling time (C) compared to the first and second
sampling times, but still remained high on the D.innoxia.
Also, it can be that D. strammonium has a high rate of transpiration and can not tolerate dry
conditions as the third sampling time was collected in June and by that time the rain has
stopped and it was hot. This may lead to most of its green intact pods to dry up, explode and
die earlier than D. innoxia. So, with the high density of buds, flowers and extreme density of
green intact capsules on D. innoxia indicated that the D. innoxia can withstand dry
conditions due to probably deeper penetrating roots that ensure enough water and nutrients
uptake also, but the rate at which its reproductive structures develop and mature might be
slower than D. strammonium.
This indicates that the two species have different flowering patterns where D. innoxia has a
continuous flowering pattern and D. strammonium reproduces, explode and dies. D.
strammonium were observed drying up with only low density of brown intact and high
density of brown exploded pods, but still hold the upright position. This was not the case
16
with D. innoxia as there were times it had all reproductive structures such as in figure 1C,
where buds, flowers, green intact, brown intact and brown exploded capsules were present
at the same time but still had greenish leaves and stems, thus indicating further potential of
continuous flowering (Macdonald et al. 1985).
significant difference between most of the reproductive structures in site one. Buds (t = 7.7
df = 264, P < 0.05), flowers (t = 8.66 df = 264, P < 0.05), green intact capsules (t = -4.18 df =
264, P < 0.05), green exploded capsules (t = 3.02 df = 264, P < 0.05) and brown exploded
capsules (t= 4.53 df = 264, P < 0.05). Except for brown intact pods where t = -1.37 P > 0.05,
and df = 264. This area site 1 (2203415.4S, 01700413.7E) was located across the tarred
road and water was blocked from flowing across. This may reduce the moisture content
necessary for plants.
It can also be that D. innoxia is highly competitive than D. strammonium because within the
vicinity of there were a high species diversity of plants such as common sunflower
(Helianthus annulus) which were abundant and various species of the family Poaceae. This
indicated that the competition between these species for resources such as water, light and
nutrients was high and only the best competitor survived. Furthermore, the area was rocky
and dry by the time of the last sampling mostly, and this might have affected also the faster
drying up of the D. strammonium and its reproductive structures. D. innoxia was subjected to
the same conditions but was able to survive. These features motivate the reason for it being
an invasive species as it can tolerate harsh environmental condition.
Human impacts could have also contributed to the drying up of D. strammonium, because
even though the habitat was located along the river, there were intensive human impacts on
the surrounding as they make use of the habitat as their dumping site. This may limit the
proper growth of the plants as they dump garbage directly on them. There was no significant
difference in the brown intact capsules of the two species (as indicated above for site 1) at
the same site as most of the green intact capsules on D. Innoxia and D. strammonium
mature and explode and straight to brown exploded.
Site two
At site 2, both the two species had buds, flowers and green intact capsules and none
developed any other capsule such as green exploded especially during the first and second
sampling time. This might be so because there was water still in the river and plants had just
then flowered immediately. In figure 2 (B), D. strammonium had an increase in density of
flowers and extreme green intact capsules. This indicates that reproductive structures on D.
17
strammonium grow so fast compared to D. innoxia, that is why the density of buds and
flowers on D. strammonium decreased in figure 2 (B), but accompanied by appearance of
green exploded and brown exploded pods as shown in figure 2 (C).
Figure 2 indicated that the plants at site 2 were still young and greener, but some of the D.
strammonium started producing green intact capsules, brown intact capsules and brown
exploded capsules as they were already drying up earlier than D. innoxia. This happened
within a short time as these plants started reproducing at very young stage, but there were
still some coming up due to the water availability as mentioned in literature (Henderson and
Wells, 1987). The soil appeared also to be rich in nutrients as it was so dark and soft
compared to the rocky soil of habitat 1 as mentioned earlier. The t-test that were again used
to compare the density of reproductive structures of D. innoxia and D. strammonium at site 2
revealed that there was no significant difference between the buds, flowers, green intact,
green exploded, and brown intact capsules, where, buds (t = 0.99 P > 0.05 and df = 338),
flowers (t = 0.23 P > 0.05 and df = 338), green intact capsules (t = 0.77 P > 0.05, df = 338),
green exploded capsules (t=2.65 P > 0.05, df = 338) and brown intact capsules (t = 1 P >
0.05 and df = 338). Except only green exploded capsules where t=3.09, df = 338 and P <
0.05. These differences results because the plants at site 2 were all young when sampling
started, and started growing and flowering at the same time and at very early age. This
could be also due to continuous presence of moisture as mentioned already. Significant
difference in the brown intact pods results because D. strammonium produces green
exploded pods earlier than D. innoxia where some green intact pods develop into brown
intact pods and other develop into brown exploded pods earlier than D. innoxia.
Relationship between crown width and reproductive structures
Table 1 indicated that there was no correlation between the reproductive structures and the
crown width of alien invasive D. innoxia and D. strammonium species. This means that an
increase in width is not accompanied by an increase in reproductive structures. With D.
innoxia there was a positive correlation between its green intact pods and crown width. This
implies that the more the crown width, the more total green pods it has, and this is not the
case to some other reproductive structures.
Ecological importance of different flowering patterns
The density of reproductive structures of the two species changes, and is also affected by
the general characteristics and structure of the habitats, that is why the t-test indicated that
18
there was a significant difference in the buds, flowers, green intact and exploded capsules
and brown exploded capsules, except only for brown intact pods at site 1, while at site 2,
there was no significant difference in the buds, green intact and brown intact capsules. It is
of an outmost importance with D. innoxia to have a continuous flowering pattern as this
gives it a chance to produce vast number of seeds and within a short period of time. This
ensures for more density of D.innoxia to germinate, and occupy large area, thus out
competing the native species as there are limited spaces for them to grow. With D.
strammonium, the fact that it reproduces once and dies encourages its seeds to be
dispersed to different distant sites faster before the next season to germinate. This is
possible as their seeds are long lived and can survive in different climatic conditions as
mentioned in literature (Henderson and Wells, 1987).
9. Conclusions
The densities of the reproductive structures on D. strammonium and D. innoxia are different
at different sites and vary at different time intervals. This allows one to conclude that the two
species have different flowering patterns where D. innoxia reproduces continuously, while D.
strammonium reproduces once, explodes dry up and dies. Therefore, it is possible for
different species that belong to the same genus such as the Datura and family such as
Solanaceae to have different flowering patterns.
19
10. References
Avery, A. G., Satina, S. and V. Rietsema, (1950). The genus Datura. The wait Ronaldo Press
Company, New York.
Boyer, H.J. and D.C. Boyer, (1998). The status of alien invasive plants in the rivers of the Namib
Naukluft Park. Directorate of Nature Conservation and Recreational resorts, Namib
Research Institute.Walvisbay, Namibia. Pg 51-57.
Bromilow, C. (2001). Problem plants of South Africa a guide to the identification and control of
more than 300 invasive plants and other weeds. Briza, pg 124- 174
Brown, C. J., Macdonald, I. A. M. and S. E. Brown, (1985). Invasive Alien organisms in South
West Africa /Namibia. South African National Scientific programme Report # 119.
Foundation of Research Development, South Africa.
Brown, J. H. and A. C. Gibson, (1983). Biogeography. The C. V. Mosby Company, London. Pg
46- 47
Bullock, J. M. Kenward, R. E. and R. S. Hails, (2001). Dispersal Ecology. Blackwell, United
Kingdom. Pg 374- 375.
th
Campell, N. A. and J. B Reece, (2005). Biology. 7 Edition. Pearson Benjamin Cummings, New
York. pg 1213.
Cowling, R. M. Pierce, S. M. and D. M. Richardson, (1997).
20
www.drugscope.org.uk/resource/drugsearch/drugsearchpages/datura.htm21/08/08
11. Appendix
Site 1 D. innoxia
Table1. raw data of site one for D. innoxia
Innoxia site 1
1st sampling
Pods
Brown
Buds
flo
grn Int
G.E
BI
E.
h(m)
w(m)
44
11
1.35
2.8
43
1.5
1.42
1.05
0.77
1.2
1.25
1.42
1.05
1.4
0.68
13
1.49
1.56
1.45
1.14
14
10
10
1.2
1.5
1.1
0.6
12
1.5
1.15
0.77
0.4
42
18
25
1.5
0.9
1.4
1.12
21
1.3
1.1
1.5
0.97
1.39
0.3
1.3
0.75
19
1.15
0.63
1.5
0.8
1.4
0.71
10
1.21
1.1
1.32
0.74
1.36
0.54
1.54
1.3
1.42
1.69
1.35
0.75
1.44
0.64
12
1.52
0.85
1.42
0.93
1.36
1.28
1.23
0.52
1.25
0.67
1.15
0.34
1.15
0.45
22
1.42
0.88
1.45
1.3
11
1.29
0.84
1.43
0.93
1.29
0.65
0.89
0.45
1.28
1.34
width
Buds
flw
grn Int
G.E
BI.
B.Ex.
height(m)
(m)
0.5
0.69
0.62
0.65
0.73
0.8
0.72
0.89
15
0.66
0.92
0.69
0.76
19
10
0.62
1.01
14
13
12
0.78
1.02
0.54
0.72
11
16
0.89
0.92
23
0.63
0.17
0.64
0.57
0.65
0.55
0.79
0.86
0.63
0.69
0.74
0.67
10
0.62
0.84
0.62
0.58
0.54
0.53
0.59
0.18
0.63
0.66
0.64
0.17
11
10
0.68
0.17
11
0.59
0.17
0.52
0.43
0.63
0.67
12
10
0.7
0.81
0.67
0.99
0.55
0.16
0.56
0.7
14
11
0.55
1.02
24
0.59
0.61
0.5
0.52
10
0.67
0.83
0.63
0.79
0.58
0.63
10
0.57
0.66
0.68
0.59
0.54
0.63
14
11
0.54
0.72
16
0.65
0.96
10
11
0.55
0.91
14
0.58
0.87
0.44
0.47
0.45
0.62
10
0.58
0.64
23
0.78
0.75
24
0.98
1.1
0.72
0.54
11
0.92
0.84
17
0.98
1.02
34
0.89
0.23
25
24
0.97
0.97
0.85
0.76
0.72
0.57
25
1.07
0.87
15
1.13
0.93
0.45
0.39
0.78
0.73
16
1.06
1.34
18
0.7
0.94
18
10
0.74
0.92
time 3
innoxia
green
green
brown
brown
width
buds
flower
intact
explode
intact
explode
height(m)
(m)
1.1
1.02
10
1.2
1.29
1.05
0.65
1.42
1.1
1.5
0.45
1.23
0.84
22
1.36
1.23
10
1.5
1.17
26
1.48
1.19
22
13
49
1.78
1.9
10
1.55
1.38
11
1.44
1.32
1.55
1.28
1.34
0.72
1.43
1.05
1.25
0.42
1.17
0.75
1.19
0.95
12
11
1.35
1.25
1.24
1.93
1.12
0.84
1.42
0.79
1.2
0.62
0.99
0.53
10
11
1.42
1.73
14
35
1.05
1.23
26
1.25
1.21
1.12
0.31
1.11
0.41
27
Site 1, D. strammonium
Table 2. raw data of D. strammonium site 1
1st time
Grn
Ex
Brwn
Buds
flower
Grn I.
br I.
57
12
21
Ex
h(m)
w(M)
1.44
1.87
1.5
1.12
1.6
1.4
11
1.8
1.3
1.2
1.14
27
1.4
1.34
15
1.36
1.22
1.29
0.82
10
1.44
1.1
1.56
0.62
13
1.55
1.32
1.47
0.65
12
1.54
1.2
1.38
0.84
1.49
1.19
1.6
1.5
28
1.69
1.1
12
1.46
0.92
1.44
0.76
1.62
0.74
13
1.67
1.16
1.45
0.99
1.12
0.75
14
1.43
1.13
29
tyme
Grn
Ex
Brwn
Buds
flwr
Grn I.
br I.
13
19
Ex
h(m)
w(M)
0.51
0.61
0.64
0.85
0.58
0.62
0.36
0.77
26
0.79
1.3
29
1.28
1.2
12
0.96
0.84
0.89
0.83
11
0.85
0.92
25
1.12
1.08
47
1.34
1.22
3rd
0
24
1.34
1.26
63
1.38
1.45
13
1.26
1.05
39
1.36
1.11
34
1.24
1.33
15
1.56
1.2
30
80
1.41
1.23
39
1.54
0.98
13
1.32
0.89
13
1.34
0.83
1.36
0.45
39
1.43
1.01
17
1.32
1.24
1.33
0.64
12
1.52
0.81
1.42
0.53
16
1.41
0.98
23
1.42
0.98
37
1.55
0.08
17
1.33
1.7
17
1.37
0.84
24
1.5
1.02
11
1.43
0.72
10
1.47
0.62
16
1.3
1.21
12
1.43
0.66
11
1.42
0.6
31
16
1.31
0.86
16
1.57
1.31
24
1.33
1.21
Site 2 D. innoxia
Table 4. raw data of D. innoxia sat site , first sampling time
Pods
width
Buds
flw
G.I
12
G.E
B.I
B. E
height(m)
(m)
0.55
0.91
0.52
0.82
0.35
0.68
15
0.58
1.07
24
0.54
1.03
13
1.1
0.06
0.3
0.6
0.35
0.64
13
0.68
0.74
23
0.9
0.78
0.75
0.8
0.6
0.7
32
0.5
0.7
10
0.66
0.68
0.34
0.51
0.53
0.47
0.34
0.33
0.41
0.49
0.36
0.41
0.27
0.26
0.39
0.45
0.35
0.34
0.33
0.31
0.32
0.32
tyme
Brwn
Buds
flw
G.I
8
11
Grn Ex
Brwn Int
Ex.
width
height(m)
(m)
0.34
0.44
0.45
0.65
0.53
0.88
33
0.33
0.41
0.36
0.53
0.44
0.78
0.41
0.52
14
0.67
0.66
0.41
0.59
10
0.54
0.85
0.37
0.48
0.46
0.49
0.33
0.49
0.33
0.34
0.51
0.59
0.44
0.49
0.47
0.63
10
0.66
0.82
0.58
0.92
0.47
0.48
0.52
0.61
0.54
0.66
0.46
0.16
0.64
0.56
34
0.53
0.72
0.51
0.51
12
10
0.79
0.93
3rd
width
Buds
flw
G.I
G.E
B.I
B. E
height(m)
(m)
0.54
0.7
0.47
0.67
0.49
0.48
28
16
30
0.73
1.38
17
14
0.82
1.27
19
18
1.02
1.43
26
10
17
0.86
1.28
13
52
0.8
1.5
12
0.72
0.92
15
26
0.93
1.42
33
0.78
1.45
35
26
0.74
1.12
12
12
0.85
1.23
14
14
0.75
1.29
0.79
0.58
0.64
0.75
0.53
0.62
0.51
0.63
0.52
0.56
0.45
0.48
0.43
0.51
10
10
0.57
0.98
0.48
0.66
10
0.48
0.18
0.59
0.72
12
10
0.73
1.13
14
16
0.67
1.11
0.76
0.79
0.53
0.64
0.54
0.68
0.64
0.65
16
0.65
0.95
36
14
20
0.86
1.2
0.41
0.78
0.66
0.73
14
0.76
0.67
12
0.67
0.72
12
0.76
0.79
0.66
0.53
0.78
0.76
0.75
0.82
20
0.89
1.06
10
0.67
0.93
0.55
0.76
0.54
0.53
11
0.65
0.85
0.47
0.49
0.55
0.64
0.69
0.59
10
0.75
0.65
0.64
0.72
11
0.76
0.74
0.59
0.6
37
12
0.63
0.78
0.75
0.5
0.63
0.67
0.65
0.58
12
0.73
0.88
12
0.72
0.88
29
0.89
1.21
11
0.94
0.88
0.64
0.52
10
0.72
0.67
12
0.72
0.65
28
0.84
1.22
0.39
0.37
0.53
0.54
15
0.74
0.85
0.53
0.55
0.58
0.59
12
0.75
0.68
0.79
0.78
0.58
0.72
0.51
0.55
38
0.54
0.87
0.74
0.91
24
0.77
0.95
10
0.73
0.78
18
0.67
1.15
14
0.74
1.05
23
0.67
1.17
24
0.94
1.13
10
0.67
0.83
0.75
0.7
0.64
0.68
0.56
0.79
0.57
0.72
0.54
0.49
0.49
0.61
22
0.79
1.07
0.55
0.78
0.54
0.69
0.38
0.55
0.5
0.81
0.45
0.59
39
0.53
0.69
0.48
0.55
0.72
0.79
0.46
0.64
0.58
0.84
0.43
0.62
0.53
0.74
31
28
0.99
1.37
0.62
0.68
0.48
0.54
0.54
0.66
0.49
0.78
0.53
0.64
0.76
0.89
14
0.69
1.05
10
19
0.65
1.03
0.55
0.76
0.59
0.73
0.54
0.89
0.68
0.67
0.65
0.77
40
0.67
0.73
10
11
28
0.85
1.43
11
15
0.74
1.08
Site 2 D. strammonium
0.53
0.84
0.25
0.38
10
42
0.9
1.4
35
52
0.85
1.06
0.76
0.67
0.79
0.62
23
100
105
1.45
Table 8. Raw data of D. strammonium second site and second sampling time
2nd
tyme
41
flower
Buds
Grn
Gre.I.
Ex
Brwn
B.I.
Ex
height
width
0.49
0.34
35
0.02
1.25
22
0.75
0.98
12
0.55
0.17
13
0.48
0.69
0.24
0.27
0.34
0.53
0.22
0.27
0.46
0.63
0.45
0.64
0.39
0.48
12
10
0.55
0.17
0.34
0.42
0.38
0.47
0.23
0.24
0.39
0.46
0.27
0.28
0.23
0.29
0.34
0.35
0.26
0.33
42
0.57
0.75
18
26
0.59
1.07
0.46
0.65
0.28
0.38
0.52
0.72
18
18
21
0.17
1.07
10
10
11
0.64
0.93
12
12
0.65
0.98
22
10
0.58
0.92
23
10
0.46
0.46
0.39
0.39
0.36
0.36
14
10
0.46
0.84
26
11
18
0.55
0.76
0.33
0.54
0.33
0.35
0.32
0.43
22
0.44
58
0.38
0.52
10
19
0.59
0.88
0.36
0.51
43
0.28
0.39
0.33
0.64
0.58
0.55
Table 9 raw data of D. strammonium second site and second sampling time
site 3 tyme
0
11
0.23
0.48
25
10
21
0.9
1.32
0.25
0.35
14
15
38
27
1.06
1.33
0.75
0.62
63
1.09
1.72
18
20
59
0.89
1.56
25
10
36
0.73
1.22
0.43
0.36
0.44
0.48
14
0.55
0.67
10
0.52
0.65
10
0.56
0.68
33
15
10
0.62
0.99
15
0.56
0.55
44
21
0.59
0.83
15
0.52
0.51
12
0.52
0.65
0.48
0.58
0.38
0.59
0.36
0.53
27
0.75
0.82
50
0.63
1.25
14
0.53
0.77
0.44
0.66
14
16
15
0.64
1.03
45
0.54
1.18
45
0.83
1.22
16
25
34
0.69
1.18
36
0.63
1.14
0.35
0.64
40
18
0.64
1.24
0.36
0.55
14
0.54
0.83
13
0.53
0.74
20
0.58
0.97
45
0.44
0.55
24
0.63
1.1
39
0.77
1.27
12
0.59
0.94
20
0.77
0.93
10
12
0.75
0.98
16
10
0.84
1.28
0.64
0.74
0.46
0.62
1.22
1.06
14
0.92
0.69
10
1.23
1.9
11
0.88
0.92
13
30
0.56
1.18
34
1.11
1.02
22
15
0.98
0.93
34
1.29
1.59
34
0.81
1.21
Data analysis
Site 1
46
Table 1
buds
Variable
Variable
Mean
4.105263
0.112782
Variance
34.91308
0.615972
Observations
133
133
Pooled Variance
17.76452
df
264
t Stat
7.724616
P(T<=t) one-tail
1.17E-13
t Critical one-tail
1.650646
P(T<=t) two-tail
2.34E-13
t Critical two-tail
1.96899
Table 2
flower
Variable 1
Variable 2
47
Mean
3.406015038
0.233082707
Variance
14.50056961
3.361927546
Observations
133
133
Pooled Variance
8.931248576
Hypothesized
Mean
Difference
df
264
t Stat
8.657959385
P(T<=t) one-tail
2.42026E-16
t Critical one-tail
1.650645911
P(T<=t) two-tail
4.84052E-16
t Critical two-tail
1.968990438
green intact
Variable 1
Variable 2
Mean
3.428571
7.849624
Variance
70.86797
77.47722
Observations
133
133
Pooled Variance
74.17259
Hypothesized
Mean
Difference
df
264
t Stat
-4.18615
48
P(T<=t) one-tail
1.93E-05
t Critical one-tail
1.650646
P(T<=t) two-tail
3.87E-05
t Critical two-tail
1.96899
green exploded
Variable 1
Variable 2
Mean
0.631579
0.142857
Variance
3.082935
0.396104
Observations
133
133
Pooled Variance
1.739519
Hypothesized
Mean
Difference
df
264
t Stat
3.021746
P(T<=t) one-tail
0.00138
t Critical one-tail
1.650646
P(T<=t) two-tail
0.00276
t Critical two-tail
1.96899
brown intact
Variable 1
Variable 2
Mean
0.090225564
0.157895
Variance
0.113009797
0.209729
Observations
133
133
49
Pooled Variance
Hypothesized
0.161369332
Mean
Difference
df
264
t Stat
-1.37369776
P(T<=t) one-tail
0.085350426
t Critical one-tail
1.650645911
P(T<=t) two-tail
0.170700852
t Critical two-tail
1.968990438
brown exploded
Variable
Variable
Mean
5.571429
0.744361
Variance
146.3528
4.661426
Observations
133
133
Pooled Variance
75.50712
Hypothesized
Mean
Difference
df
264
t Stat
4.530021
P(T<=t) one-tail
4.47E-06
t Critical one-tail
1.650646
P(T<=t) two-tail
8.94E-06
50
t Critical two-tail
1.96899
Table 1
Variable
buds
Variable 1
Mean
4.576471
3.929412
Variance
33.96157
38.62221
Observations
170
170
Pooled Variance
36.29189
df
338
t Stat
0.990258
P(T<=t) one-tail
0.161378
t Critical one-tail
1.649374
P(T<=t) two-tail
0.322756
t Critical two-tail
1.967007
flowers
Variable
Variable 1
Mean
2.964705882
2.858824
Variance
28.48395405
7.376401
51
Observations
170
Pooled Variance
17.93017751
df
338
t Stat
0.230537061
P(T<=t) one-tail
0.40890697
t Critical one-tail
1.649374277
P(T<=t) two-tail
0.81781394
t Critical two-tail
1.967007242
170
gren intact
Variable
Variable 1
Mean
7.835294
6.876471
Variance
194.4698
66.07341
Observations
170
170
Pooled Variance
130.2716
df
338
t Stat
0.774503
P(T<=t) one-tail
0.219587
t Critical one-tail
1.649374
P(T<=t) two-tail
0.439175
52
t Critical two-tail
1.967007
green explode
Variable
Variable 1
Mean
0.564706
0.105882
Variance
3.584546
1.51535
Observations
170
170
Pooled Variance
2.549948
df
338
t Stat
2.649046
P(T<=t) one-tail
0.004225
t Critical one-tail
1.649374
P(T<=t) two-tail
0.008451
t Critical two-tail
1.967007
brown intac
Variable
Variable 1
Mean
0.082352941
Variance
1.152941176
Observations
170
170
Pooled Variance
0.576470588
53
df
338
t Stat
P(T<=t) one-tail
0.159012934
t Critical one-tail
1.649374277
P(T<=t) two-tail
0.318025868
t Critical two-tail
1.967007242
Variable
brown exploded
Variable 1
Mean
1.752941
Variance
54.45931
Observations
170
170
Pooled Variance
27.22966
df
338
t Stat
3.097104
P(T<=t) one-tail
0.001059
t Critical one-tail
1.649374
P(T<=t) two-tail
0.002118
t Critical two-tail
1.967007
54