Beruflich Dokumente
Kultur Dokumente
3. CEDAR ECOLOGY
11. REFERENCES
Widdringtonia cedarbergensis (The Clanwilliam cedar)
A Synthesis of information
Initially this document was compiled to provide background information for a workshop in 2002, Cedar
conservation – the way forward, that was held at CapeNature’s premises in Porterville. It has
subsequently been updated in 2008
Thus, Widdringtonia cedarbergensis represents one of about 630 global species of conifers,
and one of nine Southern African conifer species. It is one of four species in a genus that is
endemic to Southern Africa. Further, W. cedarbergensis is an endangered species that has
been given global importance due to being short-listed by the IUCN onto a list of 43 conifer
species worthy of special conservation attention. It warrants our efforts to do what we can to
prevent its extinction.
2
Nardouw Sandstone Formation above the shale band. The only natural occurrence on the shale band
is in the east, where they grew prolifically forming the only remaining closed stand of cedars prior to
fire in 1985 (Manders 1985). They grow mainly on the eastern slopes in the northern areas, but are
not limited to any aspect elsewhere (Andrag 1977).
The climate of the Cederberg is Mediterranean with most of its annual rainfall occurring in winter, and
with hot, dry summers. The overall vegetation is fynbos comprising a complex variety of habitats and
a diverse vegetation. In the northern Cederberg area alone Taylor (1993, 1996) distinguished 26 plant
communities. He found that the main concentration of cedars occurred in an open woodland with a
mid-dense restioid understorey, and associated with protruding bedrock and cliffs. The cedar plant
community is not a well defined floristic unit, and the habitat is neither rare nor specialised, suggesting
that cedars may have been more widespread in the past. Further, none of the fynbos species typical
of this community are critically rare or narrowly endemic, suggesting that the cedar plant community
as a whole does not require special protection.
Recent studies by Higgins et al (2001) whereby the environmental preferences of the cedar were
investigated, confirmed the association of high (ca. 80%) rock cover with cedar distribution. Cedar
abundance was also associated with low vegetation cover, and with intermediate levels of solar
radiation in December.
The association of extant cedar trees with rocky areas is widely attributed to the protection thus
afforded from fire (Manders 1986a, Privett & Bond 1994, Higgins et al 2001, Fox 2003). The survival
of planted seedlings is also associated with the higher moisture regimes of rocky microhabitats
(Manders & Botha 1987, Mustart et al 1995), a factor also likely to influence its distribution. Other
suggestions that attribute the association of cedars with rocky habitats are the possibilities of directed
dispersion by baboons who take cones to rocky outcrops for eating; and the existence of underlying
water resources (Higgins et al 2001).
3. CEDAR ECOLOGY
Studies have shown poor correlation between rainfall and tree ring width of wood cellulose (surrogate
for growth) (Dunwiddie & La Marche 1980, Zucchini & Hiemstra 1983, February & Stock 1999)
implying that water is not a major limitation to the growth of the Clanwilliam cedar. It has been
suggested that the cedar has a deep root system allowing access to water irrespective of rainfall
variation (February & Stock 1998, 1999).
The Clanwilliam cedar grows in fire prone fynbos vegetation and is killed by fire. Modern thought on
the dynamics of cedar population ecology is dominated by this factor. Dependence on adequate seed
production, post-fire germination and seedling establishment of this obligate reseeding species is
critical to its persistence.
It is weakly serotinous. However, high levels of predation by beetles, baboons and rodents (Botha
1990) leave few cones remaining, making it in effect non-serotinous. The nut-like seeds take about 30
months to mature in their cones (Manders 1985, Botha 1990), with seeds being released after fire, or
after the cones are damaged. The wingless seeds drop to the ground and are poorly dispersed
(Manders 1985). The possibility of baboon-directed dispersal (Higgins et al 2001) is discussed above
(see 2).
Experimental work has shown that ground litter underneath stands of Widdringtonia cedarbergensis
suppresses germination of its seeds, hence much of the germination in the wild occurs after fire which
consumes the litter (Manders 1987a). However, inter-fire recruitment does occur, and multi-aged
cohorts are common (Midgley et al 1995, Fox 2003) as was evident in the cedar populations near the
Middelberg hut area (Mustart personal observation) which had not burned for about 100 years prior to
the 2001 fire. Fox (2003) found high inter-fire recruitment rates in two years that had had high annual
3
rainfall, postulating that in the wetter Quaternary period when cedars were more abundant, it was rain
events rather than fire that stimulated cedar recruitment.
Fynbos vegetation in the Cederberg was shown to burn at an overall frequency of 11 - 15 years
(Brown et al. 1991). However, fire frequency has accelerated in some areas in more recent times - an
example is given by Fox (2003) in her study area where 6 fires occurred in 26 years (1977 – 2003).
Individuals produce first cones after about 12 years, reaching full reproductive maturity only after 30
years (Andrag 1977), a period far longer than the average fire interval in recent times.
4
5. ATTEMPTS TO CONSERVE AND MANAGE CEDAR POPULATIONS: 1805 - 1987
i. Excessive harvesting (see 4.2 & 5.2) which has reduced and fragmented cedar populations to a
bottleneck situation from which it cannot recover.
ii. Livestock grazing which has destroyed young plants (see 4.2 & 5.3)
iii. Climate change over the past 14 000 years (Meadows & Sugden 1991) which may have led to
unfavourable environments and / or fire regimes for cedar persistence.
iv. Relatively recent human-induced global climate change of increased heat and dryness in
northwestern fynbos areas whereby cedar survival would be detrimentally impacted. This needs
documentation.
v. Inappropriate or altered fire regimes, whereby frequent fires lead to excessive death of young
plants, and intense wildfires which cause high adult mortality (Haynes & Kruger 1972, Andrag 1977,
Manders 1987b, Privett 1994a, Higgins et al 2001, Fox 2003). It is argued that prior to the influence
of man, the cedars might have existed in denser stands, with resulting suppression of understorey,
fire-prone fynbos vegetation, and less probability of frequent or intense fires (Manders 1987b). Hence
man-induced changes in the status of the cedar have led to unacceptable losses of mature trees in
natural wildfires, with subsequent small yield of seed and seedling recruitment. Trees take about 12
years to produce their first crop of seed-bearing cones, taking a further 10 years or so to build up
adequate seed reserves, thus it follows that the current trend of high frequency fires will lead to local
population extinctions (see 3).
vi. Excessive granivory and destruction of seedlings by rodents, dassies and baboons (De Hoog
1968, Andrag 1977)
vii. None of the above: the current cedar status is in a healthy condition, and has not been seriously
impacted by any of the above. Historical reports of excessive harvesting are 'romantic exaggerations”
(Higgins et al 2001). However, this study contradicts itself by finding that small alterations in
parameters do lead to cedar extinction (see 7.3). In this regard Fox’s finding that seed production had
diminished in certain areas (see 7.4), is significant.
6
7. PREDICTIONS REGARDING THE FUTURE VIABILITY OF CEDAR POPULATIONS
During the past two decades there have been several studies which have aimed to determine whether
cedar populations are capable of persistence and expansion; whether they would become extinct
regardless of conservation management strategies; and which strategies would be most beneficial for
cedar persistence. Their findings are summarised below:
7.1 Mander's transition matrix model (1987b) is based on the probability of transitions between
various stages in the life cycle of the species. Data were gathered from permanent plots between
1970 to 1984 during which time two wildfires and two prescribed burns occurred. The transition from
seed to seedling was emphasised as the key life history phase. Results indicated that populations of
Widdringtonia cedarbergensis are capable of increasing in the absence of fire, and that to conserve
existing stands, there should be intervals of 15 to 20 years between fires. However, the severe wildfire
in 1988/9 caused far greater adult mortality, and less regeneration, than had been predicted,
emphasising the difficulty of working with data from only a few fires. To quote from Manders' paper -
"Harper's (1977) observation that the study of tree population dynamics is a study of short cuts is
particularly relevant in this case, where mature trees may reach an age in excess of 400 years".
7.2 The transition matrix model of Privett & Bond (1994) incorporated mortality data collected
from prescribed and wild fires of various intensities in one prescribed and two wild fires between 1984
and 1989. Their results indicated that survival of the larger size classes was the key phase of the
cedar life cycle. Mortality of large tree sizes in hot wild fires at a frequency of 15 -20 years would lead
to unacceptable mortality and ultimately to local population extinctions. A fire interval of about 80 years
is necessary to enable populations to recover from intense, summer fires. However, late autumn/ early
winter prescribed burns which are relatively cool, at intervals of between 12 to 20 years, would enable
populations to expand.
b) In the cedar reserve (see 9.3) short-term patch burning should be administered to reduce the
chance of wild fires, and to protect existing trees. Seedlings should be planted out in fire-protected,
rocky microsites. This strengthened the basis of the then existing Cedar Restoration Programme of
CNC (see 9.3).
7.3 Higgins et al (2001) modelled the population viability of cedar populations using re-worked
demographic data that had been collected by Manders (1986a) and Privett (1994a), as well as other
simulated parameters that would influence cedar persistence. They modelled different scenarios,
many of which predict the persistence of cedars. For example, when a modal fire interval of 9 years
was used, persistence was the outcome. However, their predictions that the cedar could persist under
some conditions without management intervention was counterbalanced by their finding that relatively
small changes in key processes could push it to extinction. For example, a 5% alteration in the
7
parameters (s.a. fire interval, rock cover, completeness of burn, seed production, recruitment, growth
rate, back ground mortality) altered the outcomes dramatically, and rapid extinction of cedars was
predicted. Their conclusion was that the cedar is in a precarious situation, despite a recent survey
performed by them in 2001 incorporating 531 cedars throughout the area of their distribution. This
survey indicated a "healthy" spread of demographic stages, as well as a good seed production. Yet
their survey revealed a relative lack of the small stages (<20 cm in diameter), a phenomenon
concurring with Mander's findings (1987b).
b) Selection of sites for seedling plantings using their model of cedar distribution
c) Selection of sites for seedling planting that are favourable for enhanced stem growth rates (more
research required in this regard).
7.4 Fox (2003) used long term data accumulated by Manders (1987b), as well as currently collected
(2003) field data of two small cedar populations, in order to test the validification of the above three
models of Manders (1987b), Privett (1994a) and Higgins et al (2001). She found that high fire
frequency (six fires in the 1977 to 2003 period) had led to a faster than predicted decline in cedar
populations than that predicted by Manders and Privett, while the Higgins et al model was more
accurate. Her study showed that the rate of decline had been rapid: a ten fold reduction in population
numbers, and little recruitment over the 26 year period of study. Seed production had declined, and
apart from trees that were protected by rocks, their numbers were severely impacted. There was
evidence of changes in vegetation due to the high frequency of fires – comparative photography
suggested an increase in the annual grass component, and a decrease in shrubs. An interesting
finding was that initial population size was a determinant of subsequent population size: below a
certain threshold, population expansion did not occur, while above the threshold population size
remained stable or increased somewhat.
However, she stresses that the study should be repeated with larger population sample sizes.
ii. since seed source influences seedling quality, the best sources s.a. from Crystal Pool, Diuwelsgat
and Heuningvlei areas, should be selected. Middleberg and Welbedacht produce inferior seed.
iii. in general, it was recommended that seed from a variety of areas be used, keeping the source
recorded throughout the seedling establishment stages so that results can be monitored.
9
a) Fire management
The reserve was subjected to low intensity, "patch burning" of small areas in late autumn / early winter
at short intervals of three to four years to reduce fire loads, thereby decreasing adult mortality, as well
as to remove an allelopathic component which inhibits germination of cedar seedlings (Manders
1987a). Whereas this burning regime is not ideal for fynbos vegetation, it was argued that the
vegetation cover in such rocky terrain is too sparse to support extensive fires in any event. Further,
management found no evidence after 5 years of such a fire regime to support the suggestion that
Asteraceae species, in particular Stoebe plumosa would be favoured by these cool burns, and
eventually dominate the vegetation. However, there is some evidence to suggest that seeds of
myrmecochorous species require hot fires in order to germinate, and such species could be
disadvantaged by these cool burns. More research on these issues is required. As stated earlier, the
fynbos plant community in which the cedar occurs is widespread, and does not comprise any rare or
endemic species, thus no special protection need necessarily be applied (Taylor 1993) (see 2).
d) Volunteer participation
Between 1995 and 1999 the seedling plantings were performed solely by BotSoc's CIG/CCG
volunteers who planted out lesser quantities, about 800 per year. Care was given to the choice of
suitable microsites for plantings, following studies done by Privett and Bond (1994) and Mustart et al
(1995) which indicated that microsites shaded by rocks significantly enhanced seedling survival.
During the above period and in subsequent years, girl learners from Herschel School (CapeTown)
assisted with the difficult task of collecting mature, seed-bearing cones in March, a difficult task as
trees were often in inaccessible, rocky habitats, and the weather usually very hot.
10
e) Dismantling of the nursery, and financing for a new one
The seedling planting programme came to a halt in 2000 after the aging nursery was dismantled due
to lack of finances for its upkeep. Motivation by BotSoc's CCG in 2001 led to a donation of R27 000
by Fauna and Flora International for the erection of a new nursery. This new nursery was
subsequently established in 2003 at Matjiesrivier Nature Reserve on the eastern border of the
Cederberg Wilderness Area.
a) Nursery establishment
The mother nursery established at Matjiesriver Nature Reserve in 2003 is managed by CapeNature
and the Cederberg Conservancy for the production of cedar seedlings. A satellite nursery at
Bushmans Kloof Reserve has been used since 2002.
12
11. REFERENCES
Andrag RH 1977. studies in die Sederberge oor (i) die status van die Clanwilliam seder
(Widdringtonia cedarbergensis Marsh) (ii) buiteligontspanning. Unpublished MSc thesis. University of
Stellenbosch
Bond WJ & Thomas JC. 1998. Fitness variation and its possible causes in an endangered conifer,
Widdringtonia cedarbergensis (Cupressaceae). Unpublished document.
Botha SA 1990. Seed bank dynamics in the Clanwilliam cedar and the implications of Widdringtonia
cedarbergensis beetle triggered seed release for the survival of the species. South African Journal of
Ecology 1: 53-9
Brown PJ, Manders PT, Bands DP, Kruger FJ & Andrag RH 1991. Prescribed burning as a
conservation management practice: A case history from the Cederberg mountains, Cape Province,
South Africa. Biological Conservation 56: 133 - 150
De Hoog RJ 1968. An investigation of the influence of rodents on the failure of natural regeneration of
the cedar (Widdringtonia cedarbergensis Marsh) in the Cedarberge. Unpublished Report.
Dunwiddie PW & La Marche VC 1980. A climatically responsive tree-ring record from Widdringtonia
cedarbergensis, Cape Province, South Africa. Nature 286, 796-797.
Farjon A. & Page C.N. 1999. Conifers. Status Survey and Conservation Action Plan. IUCN, Gland,
Switzerland and Cambridge, UK.
February EC & Stock WD 1998. The relationship between ring width measures and precipitation for
Widdringtonia cedarbergensis. South African Journal of Botany 64: 213-216
February EC & Stock WD 1999. Declining trend in the 13C /12C ratio of atmospheric carbon dioxide
from tree rings of South African Widdringtonia cedarbergensis. Quarternary Research 52: 229-236
Fox S 2003. An assessment of the population status and demographic models of Widdringtonia
cedarbergensis. Unpublished Honours thesis, University of Cape Town.
Haynes & Kruger 1972. Controlled burning in Widdringtonia cedarbergensis populations: winter burns.
Unpublished report, Department of Forestry
Higgins S, February E & Skowno A. 2001. Distribution and viability of Clanwilliam cedar
(Widdringtonia cedarbergensis, Cupressaceae). Final Report to WWF-SA Table Mountain Fund
Hilton-Taylor C. 1996. Red Data List of Southern African Plants. Strelizia 4: 117 pp
Hubbard CS. 1937. Observations on the distribution and rate of growth of Clanwilliam cedar
Widdringtonia juniperoides Endl. South African Journal of Science 33: 572-86
Luckhoff HA. 1971. The Clanwilliam cedar. Its past history and present status. Mountain Club of
South Africa Journal 74: 33-39
13
Manders PT 1985. The autecology of Widdringtonia cedarbergensis in relation to its conservation
management. MSc Thesis, University of Cape Town, South Africa.
Manders PT 1986a. An assessment of the current status of the Clanwilliam cedar (Widdringtonia
cedarbergensis) and the reasons for its decline. South African Forestry Journal 139: 48-53
Manders PT 1986b. The effects of shading on nursery grown seedlings of the Clanwilliam cedar.
South African Journal of Forestry 138: 15-22
Manders PT. 1987b. A transition matrix model of the population dynamics of the Clanwilliam cedar
(Widdringtomia cedarbergensis) in natural stands subject to fire. Forest Ecology and Management 20:
171-86
Manders PT & Botha SA 1987. Experimental re-establishment of the Clanwilliam cedar Widdringtonia
cedargergensis: a preliminary study. South African Journal of Wildlife Research. 17 86-90
Manders PT, Botha SA, Bond WJ & Meadows ME 1990. The enigmatic Clanwilliam cedar. Veld &
Flora: Journal of the Botanical Society of South Africa. 76: 8-11
Meadows M & Sugden J. 1991. A vegetation history of the last 14 000 years on the Cederberg, south-
western Cape Province. South Africa Journal of Science 87: 34-43
Midgley JJ, Bond WJ & Geldenhuys CJ 1995. The Ecology of Southern African Conifers. In: (Ed.)
Enright & Hill. Ecology of Southern Conifers
Mustart PJ (editor) 1993. Proceedings of the Workshop: The Clanwilliam cedar: what is being done?
Botanical Society of South Africa FCC Report
Mustart P & Bond W. 1994. The decline of the Clanwilliam cedar - natural or man made? Africa -
Environment and Wildlife 2: 80 - 81
Mustart PJ, Makua C, Juritz J, Van der Merwe SW & Wessels N 1995. Restoration of the Clanwilliam
cedar, Widdringtonia cedarbergensis: the importance of monitoring seedlings planted in the
Cederberg, South Africa. Biological Conservation 72: 73-76.
Mustart P & Privet S. 1998. The Cederberg Interest Group. Veld & Flora: Journal of the Botanical
Society of South Africa. 84: 4-5
Privett S 1994a. Restoration of the Clanwilliam cedar, Widdringtonia cedarbergensis: a study on the
potential for fire as a management tool. Unpublished Honours dissertation. Department of Botany,
University of Cape Town
Seydack AHW 1992. Fire management options in fynbos mountain catchment areas. South African
Forestry Journal 161: 53-58
Schellevis N & Schouten J 1999. Clanwilliam cedar (Widdringtonia cedarbergensis J.A. Marsh), In:
Conifers. Status Survey and Conservation Action Plan (compiled by Farjon A. & Page C.N.) IUCN,
Gland, Switzerland and Cambridge, UK.
Smith CA. 1955. Early 19th Century records of the Clanwilliam cedar (Widdringtonia juniperoides
Endl.) Journal of the South African Forestry Association 25: 1-8
Taylor HC 1993. What plant communities do the cedars grow in? In: Mustart (ed) Proceedings of the
workshop: The Clanwilliam cedar: what is being done? The Botanical Society of South Africa FCC
Report.
Van der Spuy J & Atherstone R 2007. Cedar tree planting day. Veld & Flora: Journal of the Botanical
Society of South Africa. p 146
van der Merwe SW & Wessels N 1993. Current Management - what is being done? In: Mustart (ed)
Proceedings of the workshop: The Clanwilliam cedar: what is being done? The Botanical Society of
South Africa FCC Report.
Voget C 2009. Plant a cedar tree for the future. Veld & Flora: Journal of the Botanical Society of
South Africa. March issue.
Zucchini W & Hiemstra LVA 1983. A note on the relationship between annual rainfall and tree ring
indices for one site in South Africa. Water 9, 153-154
15