A Synthesis of information on Widdringtonia cedarbergensis (The Clanwilliam cedar

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By: P J Mustart Report of the Botanical Society of SA , 2002. Updated 2008 1. THE CLANWILLIAM CEDAR: A unique South African conifer: 2. CEDAR DISTRIBUTION AND HABITAT 3. CEDAR ECOLOGY 4. HISTORY OF THE DECLINE OF THE SPECIES 4.1 Archaeological records 4.2 Historical records 5. ATTEMPTS TO CONSERVE AND MANAGE CEDAR POPULATIONS: 1805 - 1987 5.1 Early recommendations for conservation measures 5.2 Control of harvesting 5.3 Control of grazing 5.4 Cedar plantings 5.5 Fire management 6. CURRENT CONSERVATION STATUS 6.1 The reasons for the current status of the cedar 7. PREDICTIONS REGARDING THE FUTURE VIABILITY OF CEDAR POPULATIONS 7.1 Mander's transition matrix model (1987) 7.2 Transition matrix model of Privett & Bond (1994) 7.3 Higgins et al models (2001) 7.4 Fox’s validification of the above models (2003) 8. GENETIC CONSIDERATIONS OF RE-ESTABLISHMENT 9. CEDAR MANAGEMENT: 1987 - 2000 9.1 General cedar area: 9.2 The cedar plantations: 9.3 The cedar reserve: the Cedar Restoration Programme 9.3.1 Progress achieved in the cedar reserve 9.3.2 Monitoring of seedling survival 10. CEDAR CONSERVATION AND COMMUNITY PARTICIPATION: 2000 ongoing 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

1. THE CLANWILLIAM CEDAR: A unique South African conifer: Conifers (or gymnosperms) comprise an ancient group of plants that evolved in the Mesozoic period 225 - 65 m.y.a. (Farjon & Page 1999). However competition from the a new group of plants, the evolutionary successful angiosperms, or flowering plants, which emerged 140 m.y.a. led to many conifer extinctions by the end of the Mesozoic period. Surviving conifers "retreated" to cold, high altitude regions, often where the soils are very infertile, where they escaped this angiosperm dominance. Today, most of the world's 630 conifer species occur in the northern hemisphere, and their slow growing, hard wood is an important source of our world's timber (eg. pines). In the southern hemisphere conifers are also represented in many families, including species of Podocarpaceae that occur in Southern Africa (eg. the yellowwoods Podocarpus elongatus, P. henkelii, P.latifolius and Afrocarpus falcatus) as well as in Australia and the SW Pacific islands. The only other conifers occurring in Southern Africa are found in the Cupressaceae family, namely Juniperus procera (pencil cedar in Zimbabwe and Kenya) and four species of Widdringtonia, a genus endemic to the sub-continent. These are: Widdringtonia nodiflora (mountain cypress, occurring in mountainous areas from Western Cape to Malawi), W. schwarzii (Willowmore cedar, endemic to Baviaanskloof in Eastern Cape), W. whytei (Malawi cedar, endemic to Malawi) and W. cedarbergensis (Clanwilliam cedar, endemic to Cederberg mountains, Western Cape). These Widdringtonia spp. are not true "cedars" and should, perhaps, more correctly be termed "cypresses". However they share characteristics such as durable, fragrant wood with cedars like the "cedars of Lebanon", Cedrus libani (Pinaceae), and are called as such. 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. CEDAR DISTRIBUTION AND HABITAT The Clanwilliam cedar is a high altitude tree, currently occurring within an altitude range of about 800m to 1 650 m (most abundant between 1200m and 1400m) on quartzites of the Table Mountain Group in the Cederberg mountains (Andrag 1977, Manders 1986a, Higgins et al 2001). Most cedars occur on the Peninsula Sandstone Formation, with some stands in the east and north east on the 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. HISTORY OF THE DECLINE OF THE SPECIES 4.1 Archaeological records Analyses of fossil pollen deposits suggest that the cedar did not exist in widespread, dense forests before present time (Meadows & Sugden 1991). However, it appears that it has declined steadily from 14 600 BP, and this is attributed to the climate becoming warmer and drier in the late Quaternary resulting in fire regimes too severe for the cedar, as well as to pre-colonial, human-altered fire regimes. Higgins et aI (2001) contest the evidence for the climate change hypothesis, citing Tyson & Lindsay (1992) who suggest that whereas it was cooler 15 000 years ago, there is weak evidence for it subsequently changing to drier in more recent times. In any event, the pollen analyses showed a sharp decline in deposits during the past few centuries, consistent with historical documentation of excessive harvesting during this time. 4.2 Historical records Early Cape colonists by-passed the rugged Cederberg mountains during forays to the north of Cape Town in the mid 17th Century, hence protecting the limited, but valuable cedar timber resources from exploitation until the mid 18th Century after farmers had settled along the Olifants river (Luckhoff 1971, Andrag 1977). The earliest known record of cedar trees was contained in a report by de Mist's Livestock and Agricultural Commission in 1805 which inspected the Roggeveld and Hantam area (Smith 1955). It was reported that half-castes made a living from cutting cedar timber extracted from the Cederberg mountains. They produced substantial beams (3m long, 15 cm square) and planks (30-40 cm wide, 4 cm thick) for distribution to the wider area. The cedar "forest" was recorded as being 40 km long and 3 km wide. Concern was expressed at the wasteful, uncontrolled method of exploitation, and especially at the destruction of young trees. The Commission obtained agreement from the local harvesters that they would sow seed in the mountain kloofs in order to implement restoration of cedar populations. In 1836 the British geographer, Sir James Alexander, who passed through the area, also commented on the wasteful method of harvesting, as well as the practise of burning to improve pasture by which many old trees and young plants were destroyed, and made a plea for preventing the future waste of valuable cedar timber (Hubbard 1937). Clearly, during the late 18th and much of the 19th Centuries, European settlers made great use of the durable and beautiful cedar wood, the only available tree in the Cederberg, for the timber needs of the wider region. A road was built in the Cederberg from Grootlandsvlaktes down to Welbedacht for transporting the harvested timber which supplied the requirements for fences, furniture, floors, doors, coffins and telephone poles. In 1879 over 7 000 pole trees were used for the telephone line between Piketberg and Calvinia, such trees are now virtually unobtainable (Andrag 1977).

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5. ATTEMPTS TO CONSERVE AND MANAGE CEDAR POPULATIONS: 1805 - 1987 5.1 Early recommendations for conservation measures (1805 - 1836) The recommendation of the de Mist Commission of 1805 to sow cedar seed into the species' natural habitat represents the first attempt to conserve populations of cedar trees, and the first attempt to conserve any forest species in South Africa (Smith 1955). It was the conceptual fore-runner of the late 20th Century pro-active Cedar Restoration Programme of Cape Nature Conservation (CNC) (see 9). A plea by Sir James Alexander in 1836 for conservation measures echoed these sentiments (Hubbard 1937). 5.2 Control of harvesting (1876 ongoing) In the latter part of the 19th Century various conservation measures were instigated to limit the unsustainable exploitation of cedars. A Forest Ranger was appointed in 1876 to manage the newly proclaimed Cederberg Crown Forest area. Three years later, in 1879, the forests were closed to the sale of living cedars and only dead trees were sold to recognised sawyers (Luckhoff 1971). This measure proved to be too little, too late. In 1883 the Superintendent of Woods and Forests deplored the loss of adult and young trees such that there were virtually no accessible trees of commercial value remaining. He also wrote that "the largest cedar standing is about 18 ft (5.5m) in girth and 70 ft (21,3m) in height but it is a dwarf compared to the big trees whose stumps are still standing as evidence of what size they were. These past giants must have been nearly double the girth of any now standing" (Hubbard 1937). This correlates with Sir Alexander's description half a century before of " a large tree of 36 ft (11 m) in girth, with 1 000 ft (305m) of plank sawn out of its giant arms" (Smith 1955). Of interest in this regard are the sizes of current large cedars. Higgins et al (2001) record that the three largest trees in a survey of 531 live trees were respectively 12 m ht, 5.5m circumference; 8 m ht, 5.5m circumference; and 4 m ht, 6.2 m circumference. Almost 120 years later, the largest trees remain smaller than those of earlier times. In 1967 harvesting of dead cedars was stopped, as deliberate incendiarism was suspected by prospective users (Luckhoff 1971) 5.3 Control of grazing (1891 ongoing) Grazing of seedlings, as well as burning to stimulate new growth during which process mature and young cedars were destroyed, was considered to be a factor in the decline of the species. Thus, in 1891 grazing by livestock was prohibited, but as this was difficult to implement, it was reversed in 1910. Grazing was once again stopped in 1937, except for two short periods during droughts in the 1960's and 1981. However, since the proclamation of the Cederberg as a Wilderness Area in 1973, grazing on the whole is not permitted ( Andrag 1977, Manders 1986a). 5.4 Cedar plantings for timber (1896 - 1914) In order to create a supply of readily available timber, and to alleviate pressure on natural populations, large numbers of cedars were established in about 120 ha at Middelberg and Heuningvlei between 1896 and 1914. This was achieved both by sowing seed, and by planting out of 78 000 seedlings reared at a nursery in Algeria. It was recorded that between 1900 and 1902 over 4 300 kg of seed was used, an amount that would be difficult, if not impossible, to collect today (Andrag 1977). 5.5 Fire management (early 1900's ongoing) Fire was recognised as a major force in the decline of cedar populations. Thus a variety of fire management policies were used from the early 20th Century onwards in order to preserve cedar numbers. Initially all cedar areas were protected from fire, later a four year burn cycle imposed, but was reverted to a no-fire policy a few years later. In 1972 prescribed burning was introduced to 5

prevent wild fires in summer that led to high cedar mortalities. Burning on a 12 year cycle when vegetation was moist was instigated, but was pre-empted by two wild fires in 1975 that burnt most of the cedar areas (Manders 1986a). (For current fire policy see www.capenature.org.za)

6. CURRENT CONSERVATION STATUS Widdringtonia cedarbergensis is classified as endangered (Hilton Taylor 1996), a category which is defined by species that face a very high risk of extinction in the wild in the near future. Further global conservation importance is accorded to the species in that a conifer specialist group appointed by the IUCN has placed the cedar on a "short list" of 43 conifer species according to certain characteristics such as uniqueness, extent of global distribution, and threats posed (Farjon & Page 1999). It is also stated that the species is likely to become extinct unless the restoration programme to plant out seedlings in the wild is continued (Schellevis & Schouten 1999). 6.1 The reasons for the current status of the cedar What has caused the current plight of the cedar? Cedar enthusiasts argue passionately about it (Mustart & Bond 1994). Reasons are attributed to one or a combination of the following: 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.

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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". Management recommendations from this study include: a) The application of prescribed burns in cedar areas at intervals as close to 20 years as possible, during late summer or autumn to minimise the death of adult trees (Manders et al 1990). b) Re-establishment using seeds, after wildfires and prescribed burns 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. Management recommendations from this study included: a) In the general cedar area the threat of wild fires should be minimised by breaking up the vegetation into a mosaic of different ages by relevant prescribed burning. To counter poor regeneration of cedars after late autumn/winter burns, seedlings should be planted in such areas. 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). Management recommendations from this study include: a) Targeting fire intensity by prescribed late summer / early autumn burns coupled with planting of seedlings, concurring with Privett & Bond (1994). Further agreement with Privett's recommendations is to increase patchiness of fuels in order to reduce the completeness of burn, thus preventing excessive tree mortality. 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. Management recommendations from this study include: The study supports the practice of seedling plantings, and motivates for fire protection (removal of vegetation round trees, and creation of fire breaks) in order to protect adult trees.

8. GENETIC CONSIDERATIONS IN RE-ESTABLISHMENT OF THE CEDAR Genetic studies on the Clanwilliam cedar indicate that it has not undergone a genetic bottleneck, and that factors such as inbreeding depression, are not a cause of the current decline of the species (Thomas & Bond 1997, Bond & Thomas unpublished). However genetic considerations are important for the future of the species, and should be considered in the practicalities of any restoration programme. Indications are that sparsely occurring trees are more likely to self-pollinate thus leading to higher inbreeding. In contrast, dense populations have a higher probability of cross-pollinating with consequent less inbreeding effects. Other findings were that seed and seedling fitness (w.r.t seed 8

germination, seedling size and drought survival) varied between different Cederberg populations, and this variation correlated with genetic heterozygosity. Thus, fragmentation effects which lead to loss of genetic diversity and consequent loss of fitness would need to be counteracted in attempts to restore the species. As a result of the above findings, Bond recommended the following: i. that the seedling planting programme should be aimed to create "groves" of trees, as opposed to small patches. 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. CEDAR MANAGEMENT 1987 - 2000 Pro-active management and restoration of cedar populations was instigated in 1987 by Cape Nature Conservation (CNC) (van der Merwe & Wessels 1993). In an attempt to prevent the continuing decline of cedars, the distribution range of the species was divided into three zones and managed as follows: 9.1 General cedar area: This area encompassed the present distribution range of cedars, excluding the plantations and the cedar reserve. In this area the management objective was maintenance of biodiversity, and the cedars were afforded no special management protection. The area was subjected to the same fire regime as that of the entire Cederberg Wilderness Area where an adaptive interference fire management system was applied (Seydack 1992). Late summer / autumn fire occurring at frequencies of 15 to 20 years were considered necessary for maintenance of biodiversity. 9.2 The cedar plantations: The cedar plantations which were established in the early 1900's (see 5.4) were protected from fire by the maintenance of fire breaks along their edges. 9.3 The cedar reserve: the Cedar Restoration Programme Here management was directed towards pro-active conservation of cedars. The objectives involved: a) An attempt to reduce the loss of adult trees in wild fires through manipulation of the fire regime. b) An attempt to ensure adequate recruitment through the planting out in the field of nursery-reared seedlings. 9.3.1 Progress achieved in the cedar reserve In 1987, 5252 ha (ca. 21%) of the cedar range was demarcated as a "cedar reserve" . The location was chosen because of an abundance of what was considered to be the ideal cedar habitat, i.e. cliffs and rocky outcrops between the altitudes of 1050 and 1650 m (Andrag 1977); the presence of locally extinct and depauperate populations; easy access via a network of tracks and footpaths; and the proximity of the Welbedacht nursery,

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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). b) Planting out of nursery- reared seedlings Experimental trials showed that seeds planted out in the wild have low levels of germination and survival (Manders & Botha 1989). About 3% - 6% of planted seeds germinate, and 0.3% - 6% survive the first summer. Seed losses were attributed to rodent granivory (Manders & Botha 1987). Thus it was considered more effective to rear seedlings in controlled nursery conditions at Welbedacht, before planting out in the wild. Sites selected for planting out of the nursery-reared seedlings were rocky out crops which provide shade, wind shelter, enhanced moisture and protection from fire (Andrag 1977, Manders & Botha 1987, Mustart et al 1995). Shading of seedlings also leads to enhanced seedling height (Manders 1986b), as was shown in an experimental, nursery-based experiment. Between 1987 and 1994 there were annual plantings of 7 000 to 8 000 seedlings into newly burnt sites by CNC Field Rangers. Each year about 6 ha was burnt and seedlings planted out at a density of about 1 000 per ha. c) Evaluation of the Cedar Restoration Programme (1993) A workshop was held in 1993 to evaluate the Cedar Restoration Programme, and to consider the way forward (Mustart 1993). Despite misgivings about the emphasis on single-species conservation to the possible detriment of wider fynbos biodiversity, participants unanimously endorsed the continuation of the project. It was agreed that the Clanwilliam cedar represents a charismatic species that easily attracts the interest of a wider public and can be used as a catalyst for concern over other conservation issues. One of the recommendations for the future included participation in the project by the wider public. This led to the formation of the Botanical Society's Cederberg Interest Group (CIG) in 1994 (Mustart & Privet 1998) which subsequently termed itself the Cederberg Conservation Group (CCG). 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.

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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. 9.3.2 Monitoring of seedling survival Monitoring the survival success of planted seedlings is vital in assessing the success of the overall restoration programme (Mustart 1993). This was initiated by Mustart et al (1995), and subsequently two full monitoring surveys were conducted by Privett (1994b) (Funded by WWF-SA) and Young (1998) (Funded by Vredendal Winery, and BotSoc through the CIG). Privett set up a protocol with permanent plots in the field, and a computer record system, for ongoing use by CNC. The survival of seedlings was found to be remarkably high during the first 10 years or so after planting out. Both the 1994 and 1998 monitoring surveys reported an average survival of over 60% for the 1987 - 1994 plantings by CNC when individually bagged seedlings were used. Later plantings which were done by CIG volunteers resulted in lower survival of 30% to 40%. These plantings involved the use of Styrofoam pannets in which seedlings had been germinated in small compartments. During the process of transplantation, damage to the seedling roots is unavoidable, no doubt contributing to the low survival rate. The monitoring studies have indicated that two year old nursery-reared seedlings survive and perform better than one year old ones (Privett 1994b, Mustart et al 1995, Young 1998). Seedling survival needs to be regularly monitored, in order to establish the best methodology for seedling plantings. This is particularly relevant after the recent years of drought (2005, 2006), and to establish levels of fire-related mortality. Fires have indeed occurred in the above areas where seedlings were planted, and documented monitoring of seedling survival needs to be re-established.

10. CEDAR CONSERVATION and COMMUNITY PARTICIPATION 2000 - ONGOING 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. b) Seed and seedling plantings Since 2002 Bushmans Kloof has hosted annual cedar planting events in partnership with the Botanical Society of SA and CapeNature. Initially these took the form of seed plantings for growth in Bushmans Kloof’s nursery until large enough for planting out. In addition, young saplings were planted out by guests in a grove on their reserve – while this grove does not fall within the natural distribution range of the cedar, the purpose was to promote an interest in cedar conservation by visitors to the area. Since 2005 annual “cedar days” have taken place at Heuningvlei, a small village in the northern Cederberg. At this event local communies participate in planting saplings in a grove, or woodlot, in the village, as well as helping to plant out seedlings into the adjacent Wilderness Area within the lower limit of the cedar’s natural distribution. GPS monitoring has enabled the survival of these seedlings to be monitored. Overall survival of the combined 2005 – 2007 plantings is 27% (Rika du Plessis, personal communication). 11

c) Creation of fire breaks After the extensive fires of late 2007 / early 2008 which destroyed many adult cedars, future seed production is consequently impacted. In order to protect extant seed-producing mature cedars, firebreaks have been made round cedar plantations in the Heuningvlei area, and more are planned to protect populations in the De Rif, Langkloof and Middleberg areas. d) Local community participation Local community involvement has played a major role in the above “cedar days” (v d Spuy & Atherstone 2007, Voget 2009): representatives from the Botanical Society, Bushmans Kloof Wilderness Reserve, Cederberg Conservancy, Heuningvlei community, local schools (in particular Elizabethfontein Primer, & Elandsfontein School) and the Greater Cederberg Biodiversity Corridor have participated. e) The cedar as an icon for conservation and education The plight and conservation of the charismatic cedar easily attracts the interest of people of all ages. The cedar has been used as an icon for engendering a wider interest and participation in environmental issues by local communities and school children (Voget 2009). As part of a school education programme in 2005, Kobie Hanekom conveyed the importance of, and threats to, biodiversity in the Wilderness and surrounding Conservancy areas through the central motif of the unique cedar tree. This has fulfilled the wish for it to be used in this way as expressed at the workshop in 1993 (see 9.3.1 c).

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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. Harper JL 1977. Population Biology of Plants. Academic Press, London 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 1987a. Is there allelopathic self-inhibition of generative regeneration within Widdringtonia cederbergensis stands? South African Journal of Botany 53: 408 - 410 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 1989. A note on establishment of Widdringtonia cedarbergensis (Clanwilliam cedar). Journal of Applied Ecology. 26: 571- 574 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, southwestern 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 Privett S 1994b. Widdringtonia cedarbergensis monitoring programme. Unpublished Report 14

Privett S & Bond W 1994. Restoration of the Clanwilliam cedar - summary report . Unpublished Report 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. Taylor HC 1996. Cederberg vegetation and Flora. Strelizia 3: 76 pp Thomas JC & Bond WJ 1997. Genetic variation in an endangered cedar (Widdringtonia cedarbergensis) versus two congeneric species. South African Journal of Botany 63: 133 – 140 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. Young S 1998. Widdringtonia cedarbergensis monitoring programme: An update. Unpublished report. 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

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