Reproduced by Sabinet Gateway under licence granted by the Publisher (dated 2012.) A female Winterberg cycad with each stem bearing a single cone. O pposite. The W interberg cy cad growing am ongst rocks in the E astern Cape. Photos: John Donaldson THE WINTERBERG CYCAD Surviving against the odds. by fohn Donaldson, Conservation Biology R esearch Unit, N ational B otanical Institute he mention of cycads conjures up images of primeval landscapes with spewing volcanoes and wandering dinosaurs. Although these images are not strictly correct, they present a graphic picture of a world that was; and now the volcanoes are quiet (except for the occasional hiccup) and the dinosaurs are gone and not even the reconstructive genius of Jurassic Park can bring them back. But what of the cycads? These extraordinary plants outlived the dinosaurs and, until recently, survived in relatively large numbers in parts of Central and South America, southern Africa and Australasia. Yet, today cycads are well known to many South Africans for one primary reason they are going extinct! Natural attrition has been greatly accelerated by the relent less removal of cycads from the wild and the destruction of their habitat over the past 50 - 100 years. The sad reality is that many T 36 cycads will inevitably share the fate of the dinosaurs, at least as far as their existence in nature is concerned. But before we sink into the mire of despair or simply sit back and wait for the cycads to die out, we need to be reminded that there are some success stories of cycads holding their own despite the tremendous odds stacked against them, giving us an insight into the attributes that have enabled them to survive for so long. How do plants that evolved with the seed ferns and dinosaurs cope in a world full of angiosperms and mammals and what can we learn about survival in an environment forever altered by human influences? A good example of a cycad holding its own is the Winterberg cycad, Encephalartos cycadifolius. As its common name implies, E. cycadifolius is restricted to the Winterberg mountain range of the eastern Cape where it occurs in Karroid mountain veld. Anyone whose image of cycads is of stylized, palm-like plants will probably be disappointed by E. cycadifolius. One of the ugly sisters of South African cycads, it lacks the majesty of its tall arboreal relatives and the graceful leaves of forest cycads. Instead, E. cycadifolius is characterized by an underground stem that suckers freely from the base so that a mature plant usually appears as a cluster of stems with each stem protruding only slightly above the ground and terminating in an untidy crown of short tough leaves. Everything about the plant suggests that it is adapted to survive in a harsh and uncompro mising environment and that its unattractive features are the very essence of its survival. In a recent update of cycad numbers throughout the world, it was estimated that there were only 4 800 E. cycadifolius plants left in the wild. Fortunately, this figure appears to be conservative, as in a study of cycad populations sponsored by the Mazda Wildlife Veld &■Flora June 1 9 9 5 Reproduced by Sabinet Gateway under licence granted by the Publisher (dated 2012.) Reproduced by Sabinet Gateway under licence granted by the Publisher (dated 2012.) The burnt rem ains of a W interberg cy cad after a veld-fire. Photo: John Donaldson Fund, two colleagues and I found more than 10 000 adult E. cycadifolius plants in the areas we visited. We were unable to get to all the known populations so the total figure may be somewhere between 10 000 and 20 000 plants. This places E. cycadifolius in a select group of only 6 South African species of Encephalartos estimated to have more than 10 000 individuals left in the wild. Most of the other 25 South African species have substantially fewer. Fire Fire occurs frequently in the grass lands of the Winterberg, destroying most of the above-ground parts of E. cycadifolius. All that remains is a charred stump that may some times be adorned by the rachi of the burned leaves. But far from destroying the plant, fire stimu lates growth and reproduction. A flush of new leaves is produced after a fire: the soft green foliage standing out in stark contrast to the blackened surroundings. The most spectacular response happens only a few years after a fire when all the mature plants in a burnt area reproduce simultane ously in a mass coning event that follows several years when no sexual reproduction takes place. This dramatic phenomenon, known as mast seeding, is most evident at firebreaks where all the mature plants on the burnt side will produce cones whereas those on the unburnt side will not. 38 This does not mean that all fires are good for the Winterberg cycad. Frequent fires are most likely to be detrimental as seed-germination and seedling-establishment are notoriously slow in most cycads, and the Winterberg cycad is no exception. Seeds germinate over a period of 18 months to 2 years and seedlings may become established only several years after the seeds are shed from the parent plant. If fire recurs within this critical period the seeds and seedlings are destroyed. A further point is that field studies have shown that seeds survive and germinate more readily under low shrubs, where they appear to be hoarded by rodents, than in open grassland. These shrubs may disappear when the veld is burned too often thereby removing potential germi nation sites for cycad seeds. When this happens, the seeds are stored under established adult cycads where they germinate in the pockets of soil that accumulate between the stems of the mature plant. But alas, these seedlings are doomed to die when they deplete the limited resources available in these small pockets of soil. Pollination Is cycad pollen dispersed by wind or insects? This has been a perennial question asked about cycad pollination at least since 1906 when Prof. Harold Pearson wrote about the likelihood of insect pollination in some South African cycads. The Winterberg cycad is the only South African cycad of w hich there is any doubt that insects are involved: when insects are prevented from visiting the female cones, fewer than 30% of the ovules are pollinated compared with 98% when insects are present. The insects responsible for pollination are two beetles that also have connections with cycads elsewhere in the world: one is M etacucujus encephalarti which is related to the genus Paracucujus found on cycads in Australia, and the other is an undescribed species in the family Lobariidae that is related to beetles associated with cycads in Central and North America. A point of concern in cycad conservation circles is that dependence on insect pollinators increases the likelihood of extinc tion because pollinators may die out and effectively emasculate the population. Already there is spec ulation that the absence of fertile seed in some cycad populations (e.g. M icrocycas calocom a in Cuba) is due to the disappearance of specific insect pollinators. Fortunately, this scenario does not seem to apply to the Winterberg cycad. Beetle pollinators abound in the male and female cones and the high proportion of fertile seed (98% ) indicates that reproduction is not limited by a shortage of pollinators. A further discovery that may have implications for the survival of other South African cycads, is that the two beetles involved in pollination are not restricted to the Winterberg cycad but also occur on several other cycads in South Africa. This means that it may be possible to transfer pollinators to other cycad populations where the beetles have declined or disappeared. Veld & Flora June 1995 Reproduced by Sabinet Gateway under licence granted by the Publisher (dated 2012.) Persistence From a conservation point of view, an important question is how long populations of the Winterberg cycad can survive in nature with out some sort of assistance? The immediate but superficial answer is that they w^ould survive for a very long time. A striking feature of E. cycadifolius populations is the stubborn persistence of estab lished adult plants. Each plant regularly produces new basal G erm inated cy cad seeds, sco rch ed by a veld-flre. Photo: John Donaldson suckers that may develop their own root systems so that estab lished adults can potentially live forever; individual suckers may die but the remainder of the plant lives on. In a study of several large populations, no adult mortality was recorded over a 5 year period although individual stems regularly died. The persistence of adult plants seems to be the main reason why the Winterberg cycad has survived because there are almost no seedlings or juveniles in most populations. The average plant has 6 stems which, given the slow^ growth rate of this cycad under natural conditions, means that these plants may be anything up to several hundred years old. The largest plant we found had 67 stems and who knows how old that must be! An unknown factor is how long adult plants will survive under veld-management designed for stock grazing, especially the regular occurrence of fire. The frequency of natural fires is estimated to be 8 - 15 years and it appears that E. cycadifolius is Veld S' Flora June 1995 adapted to reproduce sexually once during this period. In the other years, resources are appar ently used for growth and the production of additional suckers, or they are stored for the produc tion of cones after fire. When fires occur more frequently, the plants may continually channel all their resources into the production of cones and not into vegetative growth. This means that the old plants will become increasingly moribund without the addition of new suckers while no new plants will be added to the population because the seeds and seedlings are charred before they can become established. As with all long-lived plants, the danger is that we may miss the fact that they are declining because the effect is subtle and may take place over a long period. A study to determine the effects of veld management on cycad populations would probably be welcomed by the local farmers who jealously guard their cycad flora and eagerly await their reproductive events. The future With so many individuals of E. cycadifolius remaining in the wild, we might be tempted to say that their future is secure. But numbers are all relative and we do not need to look beyond southern Africa to find several examples of plants and animals where popula tion numbers have declined spectacularly within a few decades. Cycads themselves provide one of the best examples. The most important first step is to ensure that adult plants are not collected from the wild because the future of the Winterberg cycad depends almost entirely on the survival of existing adult plants. Fortunately, the Winterberg cycad seems to have so far escaped the excessive attentions of avaricious collectors, and local farmers certainly deserve credit for protecting these plants which all occur on private land. The inaccessibility of E. cycadifolius populations and difficulties experienced in trans planting this cycad have probably also deterred would-be collectors. On the positive side, E. cycadifolius is readily grown from seed and the demand among collectors could easily be met from propagated plants. A vast excess of fertile seed is produced during a mast year and only a small propor tion of the seeds is dispersed to sites suitable for germination - the remainder lie on the ground in their thousands where they desic cate and die. As long as dispersed seeds are left alone, the remaining seeds can probably be harvested without affecting the viability of the population in any way. In this way, farmers may be able to obtain a small return on their investment in the well-being of the Winterberg cycad. At the same time, inter ested growers and collectors can enjoy this unusual plant while the adult cycads in the wild can look forward to several more centuries in the sun. # A cknow ledgm ents My thanks to V ictor Pringle and Noel Ross for their generous ad vice and support and to the M azda W ild life Fund for its sponsor ship o f the Cycad C onservation P roject. THE CYCAD CONSERVATION PROJECT The Cycad Conservation Project was launched in 1992 by the National Botanical Institute with the aims of promoting cycad conser vation through the propagation and cultivation of threatened species in the National Botanical Gardens. A research component will also determine priorities for conserva tion, the chances of survival for different cycad populations, and the best management strategies to ensure long-term survival of wild cycad populations. As research necessitates extensive visits to natural populations, a vehicle dedicated to this purpose was provided by the Mazda Wildlife Fund. If you would like to help sponsor the programme, which has developed close links with other organizations involved in cycad conservation including the Botanical Society, or would like more information, contact Dr John Donaldson: tel (021) 762 1166, fax (021) 762 3229. Wiu'ure R:np NMl l H -■.!*». 39
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