C. quadrangularis is native to southern and eastern Africa and from Arabia to India.  As its name indicates, it bears four-sided stems and three-lobed leaves.  It tolerates a wide range of environments from full sun to shade. 

C. tuberosa, a native of Mexico.  It bears deciduous tendrils, which it sheds in severe droughts.  Its greenish, woody like tuber has interesting markings. 

However, there are about 200 species of these climbing plants, which are natives of several tropical and subtropical countries.   They are members of the Family Vitaceae, which is best known for plants of the species Vitis vinerifera, or the grape.

C. trifoliata with a wide range of distribution from southern USA to Mexico to the Caribbean  has very tuberous roots and 4 to 6 ridged warty stems .

C. discolor from Indonesia bears leaves which appear to be a bit more succulent, but not much.  It bears ovate, velvety green leaves mottled with silvery white and reddish-purple underneath    Its flowers are deep purple.

C. antarctica, native of Australia is shrubby and climbs along with tendrils. It has shiny, green leaves with brown veins and stems.   It is not very succulent at all. 

Another very leafy species is C. rhombifolia with its hairy brown branches and shiny green leaves that have fuzzy, white undersides.   It also climbs with the aid of tendrils. In the warmest parts of the U. S., several varieties can grow outside in regular soil with partial shade. 

The best planting medium to use would be equal parts of soil rich in organic matter mixed with perlite or pumice.  Repot these plants in spring and summer.

During the summer, they need a lot of water; in the winter, less is needed, unless of course we are going through a winter heat wave, which is common in South Florida.  Do not let them dry out completely-the non-succulent species should be allowed to remain a bit moist. 

In early spring, pruning can be done to thin out the plant and restore their shape by cutting back the stray vines.

Propagation of these plants from cuttings of young shoots is easy.  I just let them dry out for a day after dipping them in Root Tone and place them in a pot with very porous mix.  This should preferably be done in early spring also. 

A word of advise as to nomenclature:  do not let yourself be confused with those plants in the genus Cyphostemma, also in the Family Vitaceae.  For years this latter genus has been interchangeably used with the genus Cissus.  I am sure that you have heard of the pachycaul, Cissus jutae-well, it is NOT that but it is Cyphostemma jutae.  Be careful.

Aloe Polyphylla

by Sue Haffner
From Cactus Corner News

            Aloe polyphylla, the spiral aloe, is one of those fantastic plants that make you want to rub your eyes. Is it real? A specimen plant, showing its spiral growth pattern, is truly awe-inspiring.

            It is also one of the world’s most endangered plants. Aloe polyphylla (the name means “many leaves”) is native to basaltic mountain areas of Lesotho, a small country entirely surrounded by South Africa. Plants cling to high altitude slopes which are characterized by copious rainwater runoff and snow in the winter. This habitat is under severe pressure from farming operations, and the plant’s only pollinator, the Malachite Sunbird, is also endangered. In addition to all this, Aloe polyphylla has been over-collected for years. Travelers have mentioned seeing local people selling plants along the roadsides or in markets. These collected plants have virtually no chance of being re-established in cultivation.

            Fortunately, nurserymen have been propagating Aloe polyphylla from seed for years, and small plants are available in the trade at reasonable prices.

            The young plants resemble some other aloes, such as nobilis, except that their leaves are noticeably softer and more “watery”. (Aloe polyphylla is 95% water.) Plants must reach a diameter of at least 8”-12” before they begin to spiral-and they may spiral either left or right-and amass about 90 leaves in order to support production of the large bloomstalk.

            In our climate, Aloe polyphylla should be protected from hot temperatures as much as possible. Most growth will occur during the cooler times of spring and fall. The plant’s roots have a high oxygen requirement, and this combination of temperature and water availability will keep the leaves firm. One sign of a plant in distress is flaccid outer leaves lying on the soil surface.

            Alan Beverly is a Santa Cruz area grower who has specialized in Aloe polyphylla for years. His firm is called Ecoscape. He recommends the following container mix: to any good commercial mix add 50% by volume orchid bark or 5/16” red lava or washed pumice. Be aware that, if you use bark, it will disintegrate in two years and have to be replaced. Don’t be reluctant to put small plants into large containers, as their roots appreciate the insulation and the free root-run opportunity. In the ground, the plants will develop best in sandy loam supplemented with organic material.

            Propagation is by seed, though seed of this species is seldom available, or by stem cuttings. Aloe polyphylla offsets only rarely. An article in the May-June 1995 issue of Cactus and succulent journal described efforts to pollinate this species. It also mentioned means employed by the Huntington to get the plant to offset (e.g., cutting out the growing point.)

            Alan Beverly considers Fusarium crown rot to be the plant’s most serious disease. This is characterized by wet purple lesions on the leaves, and can be treated with Benomyl or other fungicides.

            While the cooler summers of Santa Cruz produce better growing conditions for Aloe polyphylla than does our hot Valley, you can still grow nice plants if you cater a bit to their needs.

When Bad Things Happen To Good Cacti

by Sue Haffner
From Cactus Corner News

            What’s this? Your favorite Rebutia has developed a pin-head? Your Lobivia looks like a belt is cinching its waist? What’s going on?

            It’s so disappointing to find your favorite plants producing abnormal growth. The usual reason for “pin-headedness”-etiolation-is that the plants were grown in too much shade and/or watered too much. Sometimes, though, etiolation can occur in plants that are in a well-lighted situation. One plant can etiolate, while the one next to it looks fine. The speculation is that the etiolated plant got off to a too-rapid start in the spring. Also, some genera may be more prone to this behavior than others.

            Characteristics of an etiolated plant are these: a thinner growing point, an increased distance between nodes or tubercules, a paler color of the stem, a “stretched” appearance.

            Sometimes a short, columnar cactus will have a narrow area about the middle of the stem. This “waist” ruins the appearance of the plant. It is usually assumed that this indicates a change in culture in the plant’s history. Etiolation might have begun, then normal growth resumed. Whatever the cause, cells collapsed and this constricted area will never “plump out” again. One friend had a Mammillaria with such a narrow waist that he had had to devise a kind of crutch to hold up the plant’s head, its narrow middle now too weak to hold it upright.

            Another ugly condition that can overtake your plants is a corky, brown epidermis that seems to creep up the plant from the soil line. The experts can’t agree on what causes this. Some genera, such as Notocactus, seem more susceptible to this than others. Even plants in habitat can display this condition.

            Aside from being philosophical about it, what can you do to return these misshapen plants to a more normal condition? Well, get out a sharp knife. You can declare, along with the Red Queen from “Alice in Wonderland”-Off with her head!

            Take off the etiolated top of the plant and discard it. Offsets will grow from the old stump and can be removed from it to root on their own. For a plant with a waist, the top can be taken off and put down to root. Be sure and let the cuts dry before putting the plant in a rooting medium. For good measure, also clean your knife with rubbing alcohol or a bleach solution when you move from one plant to another.

            The pictures below are from Cactus & succulent journal, March-April, 1988.

Is It a Succulent? or Is It a Cactus?

by Lou Kilbert  
Spinal Column, Michigan C&S

The word “succulent” is a general term that applies to many plants that are often totally unrelated except for in the far, far distant past! All cacti are more or less succulents - but most succulents are not cacti. In the beginner’s mind, “cactus” is often applied to any spiny plant; although, why beginners don’t then call roses “cacti” is beyond me! On
the other hand, beginners often classify any fat plant without spines as a “succulent”. This brief article attempts to clarify the issue for the beginning enthusiast.

The name “cactus” is applied to a plant family, thus the “cactus family”, just like the lily family or the rhododendron family, etc. Cacti are plants that are exclusively American. Before the European expeditions to the “New World”, there were no cacti in the “Old World”. Today, because of mankind’s intervention, cacti are found on all continents and almost every major island. Cacti are subdivided into three subfamilies: the Pereskiae, the Opuntiae and the Cereae. The Opuntiads comprise the largest subfamily of cacti. Most are easily recognized by the “Bunny Ears” growth habit that most assume. Opuntiads
are found from the tip of South America on Tierra del Fuego to the Yukon Valley of North America. The two Michigan native cacti are both Opuntiads: the Michigan Prickly Pear (Opuntia compressa) found throughout the lower peninsula wherever the soil type and microclimate are favorable and a little known, very hardy and attractive species from
the Porcupine Mountains of the upper peninsula (Opuntia fragilis). The pads on that cactus are more or less rounded to almost globular rather than having the more typical “mitten” shape. Opuntiads are identified by:1) The fact that, when in active growth, true leaves are present. These are the little green cylinders at the growing tip. They usually fall off as the pad matures. The pad (mitten) is actually a flattened stem and not a thickened leaf as beginners often assume.

2) The second characteristic that all Opuntiads have is the presence of  “glochids” or microscopic, hooked spines. These are not the easily seen large spines that many Opuntiads also have but the furry looking buttons that stud the surface of the pad. These are the things that make many enthusiasts steer clear of this subfamily. These tiny glochids seem to jump off the plant and attack you when you least expect it. To remove
them, try pressing tape against the skin and then rapidly pull the tape off. You can also apply white milk glue (Elmer’s Glue), allow it to dry and pull it off along with many of the glochids. I have even used a razor to shave off the part that sticks out of the skin; you will still get a highly irritated reddened welt or ulcer from the embedded hooks.

3) “Areoles” are a characteristic of all true cacti found in all three subfamilies. These structures produce spines and flowers, and in Opuntiads, the areole also produces the glochids and true leaves.

The second largest subfamily of cacti, the Cereae, contains the plants that many of us grow to love. Here you will find such a great variation in size, shape and general appearance that beginners have difficulty finding the relationship between them. This subfamily contains columnar plants and globular plants. In general, the plants have ribs. Ribs allow a plant to expand to absorb large quantities of water; at that time, the ribs may seem to disappear. When “starved” for water, the ribs become more prominent. Areoles are found along the outer edges of the ribs. Sometimes the ribs get further subdivided into “nipples”, as in Mammillaria; each nipple is tipped with an areole.

Also included in this subfamily are the Jungle Cacti: Christmas, Thanksgiving, Easter, Rice and Orchid cacti. Except in the case of a few Rhipsalis (Rice Cacti), the relationship of these cacti to others in the group is not easy to see. If you ever grow one from seed and sometimes when they are grown under stress (low light and high humidity), tiny stems resembling miniature columnar cacti will be produced and then you may be able to see that these Jungle Cacti are closely allied with the others in the subfamily.

The third subfamily, Pereskiae, is not usually grown by hobbyists, but may be seen in botanical gardens. These are “primitive” members of the family that look like thorny roses that grew large enough to be called trees. The true leaves are large and waxy looking. The flowers look like single roses. However, areoles are present and produce the leaves, spines and flowers. And no, cacti did not originate in prehistory from the rose family. Pereskias are grown throughout the tropics as fruit trees. The fruit has the taste of a not-so-tart lemon.

Succulents are plants that take up or absorb water very quickly and store it in special water storage cells and organs. Succulents lose water very, very slowly 1) because they have reduced their surface area to a minimum, 2) because they cover their surface with waxy, water-resistant coatings and 3) because they greatly reduce the number of stoma or breathing pores to a minimum. Notice this descriptive definition doesn’t say anything about spiny-ness because succulents may or may not have spiny appendages. Also notice that the definition is so general that it applies to many totally unrelated plants in many
different plant families.

Cultivation of succulents is based on this definition. Succulents can take up too much water too quickly, mostly in cultivation. In nature, abundant water just is not available to the plants. After taking up all this excess water, because they lose water very, very slowly, they suffer and may even die! They can’t get rid of excess water. They have
reduced stoma, so water can’t be excreted through respiration. They have thick coatings on their stems and leaves that won’t allow excess water to escape. Thus, one waters carefully to prevent excess uptake. Christmas cactus suffer from “silver-skin disease”; this is caused by over-watering; the excess water gets trapped between the living green
tissue and the thick outer skin. When the water is eventually eliminated, the skin appears silvery because now the layer between the skin and the living tissue is filled with air. The shape with the greatest volume and minimum surface area is a sphere. Many families have evolved species that are more or less spherical in shape as an ideal adaptation to life in dry climates. Look at pictures of Euphorbia obesa and Astrophytum asterias; two totally unrelated plants with an almost identical appearance!

All cacti are succulents. Other succulents often resemble cacti as a result of parallel evolution. The family of plants most often mistaken for cacti are the Euphorbias of Africa. The Euphorbia family is distributed worldwide in contrast to the cacti that come exclusively from the Americas. Most of the Euphorbias outside Africa are shrubs and
sub-shrubs and of little interest to collectors of succulents. If spines are present, Euphorbias are easy to distinguish from cacti. Euphorbia spines are branched, often with just two “horns”, but sometimes having multiple branch points. Cactus spines are never branched; although, some are delicately feathered. Also unlike cacti, the twin horned spines of Euphorbias are mounted on a horny plate; this base plate often runs the
full length of the rib, but may be just a little shield at the base of the spine.

Euphorbias are filled with a milky sap that runs like white blood from their wounds. The sap eventually clots into a rubbery latex. Euphorbias were once investigated as a possible source of latex for the manufacture of “rubber”, but that effort was abandoned. Caution! The milky sap of Euphorbias is very acidic and can burn the eyes, mouth and sensitive
skin. Many people develop an allergic reaction to the latex as well. Some cacti, especially certain Mammillarias, also produce a milky latex; therefore, this is not a foolproof distinguishing characteristic.

Along with Euphorbias, the most often cultivated succulents come from the Lily and the Crassula or Jade Tree families. Even though Agaves and Haworthias look totally different from the vast majority of cactus, we still get people who ask, “Does that belong to the cactus family?” In the general public’s mind “cactus” is indistinctly defined and the word “succulent” is almost never used, even though that is the simple, broadly encompassing term. I guess it’s up to us to keep explaining to people the meaning of these words.

Quizzical Plants: Monstrose and Crested Succulents
by L. M. Moe
The Cactus Patch, Bakersfield, CA

To understand what causes monstrose and crested succulents, a brief look at how plants grow is necessary. All plants grow in length by increasing the number of cells at the tips of shoots and roots. These regions of active cell division are called apical meristems and this growth is called primary growth. Some plants, most notably the monocots (grasses, lilies, aloes), have meristems (called intercalary meristems) at leaf bases that cause leaf elongation. (This is why grass leaves continue to grow after they have been mowed). In woody plants there are additional meristems called lateral meristems that produce secondary growth. The most important lateral meristem is the vascular cambium, which produces wood and bark. Monstrose and crested growth only involves primary growth.

Cell divisions in the apical and intercalary meristems are highly regulated and finely tuned in each kind of plant, resulting in distinctive stem and leaf shapes. For example, very rapid apical growth with a suppression of branching (called apical dominance) results in long, slender stems whereas, slower growth results in "fatter" stems. Stems without apical dominance are highly branched. In general, cacti have apical dominance with slow apical growth and are "fat" (and the cells filled with water, making them succulent). So, in cacti faster growth results in columnar cacti and slower growth results in barrel cacti. Occasionally something called a growth mutation happens that "messes up" these regulated and coordinated cell divisions in the primary meristems. The causes of these mutations range from injury to bacterial or viral diseases. The three most common types of growth mutation are crests, monstrose growth and variegation.

In crests the growth mutation changes the shape of the apical meristem. Instead of a single growth tip the area of active cell growth becomes a line, resulting in fan-like or crested growth (see photos of crested Euphorbia resinifera and crested Trichocereus).
Euphorbia (117kB)
Trichocereus (84kB)

In monstrose growth, the local apical dominance is lost and every growth tip tries to grow as if it were the dominant point, resulting in a "knobby" or "lumpy, jumbled" growth.

Variegation can be a topic for a future newsletter, if there is interest. Please let me know.

Cresting and monstrose growth is not unique to succulent plants. Crests are found in many genera of non-succulent plants, including conifers and many common garden plants.

Crests and monstrose plants are grown the same as normal plants of the same species except that crests and monstrose plants tend to be more sensitive. This is one of the many reasons they are often grown as grafts. Monstrose and crested plants flower and produce seed just as other plants do. However the growth mutations are not generally transmitted by seed so the best way to propagate these plants is by cuttings.

In Defense of Dactylopsis
By Matthew R. Opel
Dept. Ecology and Evolutionary Biology, University of Connecticut, Storrs, CT 06269
email: matthew.opel (at) uconn.edu
San Francisco Succulent and Cactus Society Newsletter

For connoisseurs of the succulent Aizoaceae, or "mesembs," there are several compellingly odd, but difficult to cultivate species and genera that have taken on an almost mythical quality. These vegetable Maltese Falcons include Muiria hortenseae, Conophytum burgeri and Didymaotus lapidiformis, but surely the strangest and most horticulturally recalcitrant of the bunch is Dactylopsis digitata, the finger mesemb. So, it came as something of a shock to the community of succulent-plant enthusiasts when Dactylopsis was subsumed into Phyllobolus (Gerbaulet, 1997). Many of us never did quite get around to changing our labels to Phyllobolus digitatus, though, and in this article I will defend the continued use of Dactylopsis, on grounds other than that of simple inertia.

Before getting into my defense of Dactylopsis, I should summarize the arguments that were used to justify the transfer to Phyllobolus. The reasoning was based on the discipline of cladistics, which is a method of discovering relationships among organisms, and reconstructing evolutionary trees (cladograms). In cladistics, species are grouped together if they have synapomorphies, which are shared derived characteristics not present in other, putatively related groups. One goal of a cladistic analysis is to find "monophyletic" groups – groups of species that include all of the descendants of a common ancestor, and exclude representatives from all other evolutionary lineages. It is the overwhelming consensus among modern biologists that cladistics is the best way to determine the evolutionary history of living things, and that classification schemes should only recognize taxa that are monophyletic.

No-one, including Gerbaulet, contests the idea that Dactylopsis is a monophyletic entity. Depending upon whom you talk to, the group contains either two species (D. digitata, the large-leaved southern form, and D. littlewoodii, the small-leaved northern form)(van Jaarsveld and Pienaar, 2000), or a single, variable species (Gerbaulet, 2001). The two forms are mainly distinguished by size, and share synapomorphies not seen in any other genus in their subfamily (the Mesembryanthemoideae): large, hyper-succulent cylindrical leaves, and tiny white flowers with filamentous petals that tend to keep their form as they dry out. The reason why Dactylopsis was sunk by Gerbaulet is that she claimed that it had other synapomorphies in common with Phyllobolus, in particular the dwarf, tuberous relatives of P. resurgens. Although Gerbaulet did not present a formal cladistic analysis, she contended that Dactylopsis was nested within the evolutionary branch (clade) containing P. resurgens, and thus that Phyllobolus would not be a monophyletic genus if Dactylopsis was maintained as a separate entity (Gerbaulet, 1997).

Based upon gross morphology, it is difficult to see what characters could possibly link Dactylopsis to Phyllobolus: the succulent, persistent green stems, smooth seeds, and smooth, waxy leaves of Dactylopsis contrast strongly with the ephemeral stems, rough seeds and warty, bladder-cell covered leaves of the P. resurgens group. The strength of Gerbaulet’s (1997) argument lies instead with anatomical characteristics. The stems and roots of Dactylopsis, and the tubers of geophytic Phyllobolus species, thicken via the production of successive rings of vascular tissue that consists mostly of spongy, water-storing parenchyma cells. These organs also become covered by a well-developed periderm (an outer layer of protective cork cells) in both genera. However, the status of these characters as synapomorphies unique to these two genera is dubious. All perennial Mesembryanthemoideae that I have examined thicken their stems and roots via successive cambia giving rise to concentric rings of vascular tissue, which tends to be mostly parenchyma in organs that do not need to support themselves mechanically, such as the stems of a cushion plant like Dactylopsis or the subterranean tubers of Phyllobolus. A periderm, too, seems to be developed to a greater or lesser extent in all of the genera of perennial mesembs that show secondary thickening growth in roots or stems. I would suggest that the anatomical characters used to sink Dactylopsis into Phyllobolus are not synapomorphies of these genera at all, but ancestral characters (symplesiomorphies) present in a wide variety of genera in the subfamily, and thus uninformative for determining evolutionary relationships.

If the link between Dactylopsis and Phyllobolus is tenuous, what are the closest relatives (sister groups) to Dactylopsis? The extreme morphological reduction of dwarf succulents like Dactylopsis can make the task of identifying synapomorphies difficult. However, I have found several characters that indicate a strong link to Aspazoma, an obscure, monotypic genus of small shrubs (Opel, 2002). A relationship between Dactylopsis and Aspazoma was first suggested almost half a century ago by Schwantes (1957), who placed both genera in the same subtribe.

The most suggestive characteristics linking Dactylopsis and Aspazoma involve the structure of the leaves. In both genera, the epidermal bladder cells that are present in most Mesembryanthemoideae have been lost, and the leaves are quite smooth and waxy. Both genera have sheathing leaf bases that surround the stem, a character that does not occur in any other genus in the subfamily. Moreover, the function of the sheathing leaf bases is the same in both genera: the leaves dry up to form a protective tunic that covers the succulent stems during the summer dormant period. This mode of dormancy is unique in the family Aizoaceae, though many genera (e.g. Conophytum) show an analogous strategy, in which succulent leaves (rather than stems) are protected by a papery tunic of old leaves during the dry season.

Another character that suggests that Dactylopsis is related to Aspazoma, and not Phyllobolus, is their succulent stems, which have a layer of green chlorenchyma in their cortex, and prominent bladder cells on their epidermis. The stems remain succulent for two growing seasons, and then lose their green cortex and develop a periderm. Nothing similar is known from Phyllobolus, though the same type of stem morphology and development occurs in the genera that are commonly considered to be related to Aspazoma: Psilocaulon and Brownanthus (secondary growth in the stems of these shrubby genera is woody, consisting mainly of fibers and tracheids, whereas the cambia of Dactylopsis mostly produce soft parenchyma). In addition, Dactylopsis and Aspazoma both have smooth-coated seeds, while seed coats in Phyllobolus tend to be rough. In terms of overall shape, the elongated pyriform seeds of Dactylopsis have little in common with either the robust, D-shaped seeds of Phyllobolus or the large, flattened seeds of Aspazoma.

Given the available morphological evidence, it seems most reasonable to maintain Dactylopsis as a separate genus, closely related to Aspazoma, but easily distinguished by its extreme succulence and unusual floral structure. Further research is required to determine how the hypothesized Dactylopsis/Aspazoma clade falls out with respect to other allied genera, such as Brownanthus, and it is probably safe to assume that more rearrangements will need to be made before the taxonomy of this group settles into a stable state.

Gerbaulet, M. 1997. Revision of the genus Phyllobolus N.E.Br. (Aizoaceae). Bot. Jahrb. Syst. 119: 145-211.

Gerbaulet, M. 2001. Phyllobolus. In: H.E.K. Hartmann, ed. Illustrated Handbook of Succulent Plants: Aizoaceae F-Z. Springer-Verlag, Berlin.

Opel, M.R. 2002. Aspazoma in the veld and under glass. Mesemb Study Group Bulletin 17: 59-60.

Schwantes, G. 1957. Flowering Stones and Mid-day Flowers. Benn, London.

van Jaarsveld, E.J. and de Villiers Pienaar, U. 2000. Vygies: Gems of the Veld. Cactus & Co. Libri, Venegono, Italy.