learning about biodiversity Liliput and Goliath:

Veld & Flora FACTSHEET SUCCULENTS

Temperature

Coping with adversity

Succulent plant diversity in South Africas low rainfall regions is exceptionally rich and fascinating. Spanning many different families, they vary from miniature tufted plants such as Conophytum to the big and bold Baobab, the worlds largest succulent. Succulents are highly specialized and evolutionarily advanced in comparison to plants from high rainfall regions. Lets explore some of the ways they cope in their low rainfall ecosystems. An ecosystem is made up of living organisms that interact with each other and their environment. The biotic components are the living organisms such as plants, animals and micro-organisms. Abiotic components of an ecosystem are the nonliving components that affect plants and animals. They include temperature, water, light and soil as well as altitude and aspect.

Red and green:

Light

Why are so many of the dwarf succulents confined to the winter rainfall areas and the arborescent (tree-like) species to the eastern and northern parts of South Africa? Here climate dictates: a short, cool, moist winter with long, dry, hot summers often results in dwarfism, whilst warm, subtropical, moist summers encourage plants to grow gigantic. Low temperatures during winter demand a dwarf growth, close to the ground, where they can make use of the warmth of the soil. During summers they go into their resting phase. The dwarf mesembs such as Conophytum, recycle their moisture from the old leaf pair to the younger leaf pair while retaining the old, dry leaf pair as a protective covering for the young leaves. Their highly advanced local seed dispersal ensures they remain within their habitat which is often on quartz gravel hills. At the other extreme, high temperatures often result in large barrelshaped succulent stemmed plants, such as the Bottle Tree (Pachypodium lealii), Baobab (Adansonia) and many of the taller Euphorbia species. Growth is cylindrical and tall, getting away from the very hot ground. The Richtersveld with its winter rainfall, covers a much smaller area (almost four times smaller) than the Kaokoveld of Namibia with its subtropical summer rain, but the latter is much poorer in species. Dwarfism allows for specialist adaptation, and the Land of Liliput can thus fit in many more species than the giants in the Land of Goliath. The dwarf succulents are very popular in succulent collections as they take up less space, and many of them can be grown indoors.

Like all green plants succulents have to trap carbon from the atmosphere for their normal food production and growth. Many succulents turn a reddish colour during dry conditions. This is the result of a pigment, anthocyanin, which is the plants solution to slowing down photosynthesis and thus dimming the suns bright rays. Succulents thus put their foot on the brakes when moisture becomes scarce. This reddish colour is not confined to succulents, and you can often see it in young leaves (for protecting the young, vulnerable tissue) or in deciduous trees and fruit which ripens (a sign that the fruits are ripe). Many succulents keep their breathing pores closed during dry, sunny conditions, only opening them during the night. How do they manage to photosynthesize, which requires CO2? During the night, the plants accumulate organic acids to which carbon is bound, which during the day, are broken down again and the CO2 released making it available without having to loose moisture via their breathing pores. This is known as Crassulacean Acid Metabolism (CAM). Not all succulents grow in full sun. Most of the smaller species grow on south-facing slopes or below Karoo shrubs that provide protection from the sun.

Avoiding predation:

Animals

Ingenious ways of storing moisture:

Water

Why are succulents leaves and stems often cylindrical or round in shape? This is the best surface to volume water storage ratio! We make use of water tanks which are cylindrical in shape for storing water. Apart from succulence many have features for coping with seasonally dry conditions. Botterboom (Tylecodon) forsake their leaves for the long dry summers, relying on their green photosyntheticactive succulent stems. Kobas (Cyphostemma) illustrated in the centre of this poster, follows the same strategy but loses its leaves during the winter. (A good example of convergence where two different plants from different families follow a similar strategy.) Most mesembs have terete leaves (cylindrical or slightly tapering, without ridges) orientated towards the sun, thus avoiding the full blast of the suns rays. All plants transpire to keep cool through their small breathing pores (stomata). How do succulents avoid too much water loss which could result in desiccation? Some succulents have a grey or whitish colour which reflects sunlight, others have a dense, whitish, hairy skin (epidermis) such as Senecio haworthii. The Giant Iceplant (Mesembryanthemum barklyi) illustrated on the left is covered with reflective glittering bladder cells.

ABOVE: Xing Quan from the Beijing University Botanical Gardens up a Kobas (Cyphostemma currorii) growing in Omavanda in Namibia.

Putting down roots:

Soil

Succulents can grow in shallow soils, even on bedrock, where there is little competition from other plants. Most plants simply cannot survive when the top layer of the soil dries out but succulents, by virtue of their succulent nature which allows them to survive until the next rain falls, can. In the Karoo with its low rainfall, many of the shrubs have deep roots which draw up moisture, whilst the succulents survive alongside them with their shallow roots and fleshy, moisture storing nature. Many succulents grow in clayey soil too, as their shallow roots enable them to cope with the low oxygen content of clay.

The high water content of succulent plants makes them vulnerable to herbivores, particularly the larger mammals. There are various ways they avoid predation. The big and bold succulent plants such as Aloe ferox and most euphorbias are spiny, and any animal would think twice before taking them on. They also have another trick up their sleeve chemicals. Aloes are extremely bitter, and euphorbias have a with milky latex that can damage eyes or burn skin. Another clever strategy some follow is to blend in with their background (camouflage). This group is usually small and humble; some resemble stones (stone plants such as Lithops and Pleiospilos), others simply have mottled green leaves (Gasteria and Senecio articulatus) which grow below shrubs and are thus difficult to see. There is also a large group of dwarf succulents that rely on other plants for their defence. They hide below spiny shrubs (nurse plants) and are thus well adapted to grow in shade which is why they make such good indoor plants. Some, like the large and very palatable Spekboom (Portulacaria afra) however, are quite without any defence. They make use of a different strategy altogether passive resistance! Any bit of vegetative material that falls to the ground when broken off the parent plant, immediately roots and forms a new plant. Karkei (Crassula ovata), Klein-karkai (C. tetragona), Ox-tongue (Gasteria), Adromischus and many others have also evolved this strategy, turning predation into procreation.

Read more
Discover more about succulents in these articles in back issues of Veld & Flora.

Charting uncertainty: Global climate change and its implications for our flora by Guy Midgley et al, vol. 88(2) June 2002 on page 70. Cliff hangers: What defines a cliff dwelling succulent plant? by Ernst van

Jaarsveld, vol. 88(4) Dec. 2002 on page 154. Desert grapes: An epeditition to the remote reaches of the southern Namib by Ernst van Jaarsveld, vol. 94(2) June 2008 on p. 82. Fanfare in the fynbos: Aloe plicatilis, a unique Western Cape tree aloe by Stephen

Cousins, vol. 96(4) Dec. 2010 on p. 164. Fog and dew in the Succulent Karoo: An indispensible source of water for arid Succulent Karoo shrubs by Ignatious Matimati et al, vol. 96(3) Sept. 2010 on p. 140. Kleinduimpie Grass: the only succulent

grass in the world by Ernst van Jaarsveld, vol. 95(1) March 2009 on page 19. The quarzite ridges of Gauteng by Michle Pfab, vol. 88(2) June 2002 on p. 56. The remarkable Kaoko Klipblom by Ernst van Jaarsveld, vol. 93(1) March 2007 on p. 42.

Text and photographs by Ernst van Jaarsveld and Caroline Voget. Download these articles at http://LABpages. blogspot.com.

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VELD&FLORA | MARCH 2012

MARCH 2012

| VELD&FLORA

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