Cyphostemma by Sue Haffner
From To the Point - Fresno
Forest or Jungle Cacti by Deborah Wisniewska-Jones
From the Cactus Factus - Toronto November 1999
Fat Plants of South America by Carl Frederick (San Anselmo, Ca.)
Cacti of Brazil By Tom Steppe (Alameda, Ca.) Growing the Semps by Lucille Tobiassen (San Francisco, Ca.)
From the San Francisco Cactus and Succulent newsletter
Gasteria by Tom Glavich
From the Communique - San Gabriel Valley Cactus and Succulent Society
Sansevierias by Dennis Kucera
From Open Gates, Gates Cactus And Succulent Society
January Cactus Of The Month - Mammillaria Bocasana - "Fred" by David Thomas
The Northern Cactophile - A poem by Jim Alger
How Cold Is Cold? by Roy "Frosty" Paramore
The Cactus Wren by Phyllis Mceuen
From the Kaktos Komments - Houston
The Mesembs by Lou Kilbert
Notocactus by Lou Kilbert
From the Spinal Column - Michigan
Haworthias by Pam Schnebelen
Copiapoa by Jonathan Watt
Turbinicarpus by Jonathan Watt
Cephalocereus senilis by Jonathan Watt
Cephalocereus senilis by Jonathan Watt
Agave by Jonathan Watt
From St. Louis - Henry Shaw C & S
Travels with Rob by Rob Skillin
From The Cactus Patch
How Cactus Made It To The Americas by Stan Yalof
From Cactus Courier (January) - Palomar C & S
To the Point - Fresno
by Sue Haffner
Formerly included in the genus Cissus, the genus Cyphostemma contains some of the most desirable and attractive of all caudiciform plants. In their nataive habitat, these extraordinary pachycaul members of the Vitaceae are rare and long-lived plants. Cyphostemma is distributed throughout the desert habitats of eastern and southern Africa, and some are endemic to Madagascar.
The species most commonly found in cultivation is C. juttae, sometimes found with its old label, Cissus juttae. It is outstanding for its much thickened, massive trunk which can reach 3 feet or more in height, often producing several stout branches. This inflated trunk is almost wholly composed of succulent tissue and is protected by a distinctive and ornamental corky rind or periderm which peels, flakes, and rolls away so that the plant appears to be shedding its skin.
The large, fleshy leaves of C. juttae are also striking, borne in terminal clusters, often elevated on short, succulent stems emerging from tips of the tapering trunks. They are a waxy, glaucous green, tinged with mauve when young; oval, with an irregular, deeply serrated margin and prominent veins. They tend to display white latex droplets on the reverse side. The flowers appear in tall, fleshy peduncles elevated above the foliage. Conspicuous clusters of red, grape-like fruit are produced, each containing one seed.
C. currori is the largest species of the genus, with a spectacular pachycaul stem. The similar C. uter is shorter, with a much-branched crown. Cyphostemma laza is a tree-like Madagascan species with a narrowly conical trunk topped with sprawling, vine-like stems.
Aged specimens of this genus are spectacular sights in botanical gardens and greenhouses, but most of them are not difficult to grow. Young plants are readily available from cactus and succulent dealers. In the wild, their leaves are shed during the dry season and in cultivation, they fall as winter approaches. All pachycaul succulent species of Cyphostemma are sensitive to excess moisture and should be kept on the dry side. One local grower used to plant her cyphos out in the flowerbed in spring and dig up the plants in winter. Last year, she decided that was too much work. When especially low temperatures were forecast, she covered her now-leafless plants with paper grocery bags (lined with plastic bags), and the plants did just fine.
Cactus Factus - Toronto November 1999
Forest or Jungle Cacti
by Deborah Wisniewska-Jones
Epiphyllum -- A genus of 16 species of mostly epiphytic cactus native to tropical America. They have numerous branches formed of short, flattened, bright green joints, and they have large, showy, often fragrant flowers. Commonly known as orchid cactus or pond-lily cactus. Epiphyte -- In botany, a plant that grows on another but does not derive its nourishment from it, such as many ferns, orchids and bromeliads. Epiphytic -- Pertaining to or having the nature of an epiphyte.
The first record of these plants was in 1753, cuttings were brought back to Europe by the early navigators to South and Central America. In 1812 Englishman Adrian H.. Haworth (the same for whom the Haworthia genus was named) first described 'Epiphyllum phyllanthus' and 'Epiphyllum' became a valid name for a Genus. This name 'Epiphyllum' took the place of 'Cactus Phyllanthus', which was the name originally given by Linnaeus. In 1819, Haworth discovered a new type of Epiphyte, which he called 'Epiphyllum truncatum'. In 1831 Germany, Link, coined the name 'Phyllocactus' ('Phyllum' meaning a leaf), using it instead of 'Epiphyllum' to describe all epiphytes, but it wasn't used by most people as they thought 'Epiphyllum' and 'Phyllocactus' were synonymous, and preferred 'Epiphyllum'. But he kept the name 'Epiphyllum truncatum' for Haworth's plant in 1819. This nomenclature stood until 1923 when Britton and Rose took the name 'Epiphyllum' for the 12 (or 16) original species. Then came Karl Schumann who brought in the name 'Zygocactus' which applies to the 'Christmas cactus' Zygocactus truncatus.
The earliest hybrids recorded were made by Jenkinson and Smith in 1830 England, the Germans and French were the next. The original crosses were made with Heliocereus specious (hee-lee-oh-sehr-ee-uhs) and Nopalxochia (noh-puhlks-oh-shee-uh) phyllanthoides (feh-lahn-thuh-oids). In 1840 Epiphyllum crenatum which flowers on the tips of it stems was brought to France and crossed with Heliocereus specious, many new flowers from light yellowish-white and rose shade to orange and deep amber were obtained. In 1890 Johannes Nicolai at his nursery in Dresden hybridized Schlumbergera, Zygocactus and Rhipsalis and introduced 300 new 'Epiphyllum hybrids. But due to a shortage of coal during World War 1 he lost all his plants froze.
In 1930 H. M. Wegener of L.A. California imported Epiphyllum hybrids into America, he built up a collection of several hundreds. Many more Americans realized that a lot more could be done to produce more hybrids in this warm climate than in Europe and slowly the hybridization work shifted back to America, but this time California. The Epiphyllum Society of America was founded in 1940 and it has established the fact that one of the original homes of these plants was along the western coastline of the USA
Most of the blooms of the 16 true 'Epiphyllum' species are fragrant and all have white flowers, through some of the outer petals have tinges of yellow, cream and strawcolour. E. cooperi flower from the base of the plant, the buds develop slowly at first and may take up to 10 weeks to open. These plants that came from the dense tropical forests of Central and So. Amer. live in humid jungle conditions. They live high in the crotches of trees, in pockets of humus, getting partial sun or shade under the swaying branches of the trees. Their branches are jointed with areoles in the sunken crenations (kree-nayt-shuns) of the edges of the stem. The main stems are usually round and woody at the base and then become flat or triangular, (both flat and triangular stems may grow on the same plant), some branches start out triangular and become flat or vice-versa. Epiphyllums don't have any true leaves and have a wax-like outer skin to prevent evaporation. Like any other cacti they expand and contract depending on the amount of water around. Branches may be up to 15 ft long and 8 inches across, but most are about 3 inches across. During down pours, rain is quickly absorbed and stored in the branches and stems, the roots retain enough moisture to keep the fine roots from drying out. They do not spread their roots like normal cacti do and their roots live in a very small confined area. They do have air roots and these feed the joints that are the furthest away from the roots. Living in the trees as they do and only getting sun from the swaying branches they remain in sufficient shade to keep them from drying out. Their roots are anchored in the humus that is caught in the fissures of the bark, their roots may be short and fine but are strong enough to support their branches which may hang down for 10 or 15 feet. The humidity may be over 90% in this dark atmosphere but as they grow high up, well above the wet ground, they have good drainage, keeping them damp but not water-logged.
What we call epiphyllum today are actually hybrids of the epiphytic cacti species native to the jungles of So. and Central Amer. and Mexico. The name Epiphyllum, epi means "upon" and phyllum means "leaf", because they produced flowers on their leaf-like stems. But they have no leaves, these are actually thickened stems or branches, most are flat but some grown in a triangular shape. Epiphyllums are not covered with spines, but have hair bristles or tiny spines in the areolar, some more than others.
Note: 'Night Blooming Cereus' is not a cereus at all but Epiphyllum Oxypetalum. The flowers of a Cereus resemble the Epiphyllum Oxypetalum. While the bud is forming, it grows pointing downward, a few days before it blooms the bud starts to point upward. The bud opens around 10:00 p.m. and dies around 8:00 a.m. the next morning.
Other Epiphytic Species:
(The following was taken directly from Myron Kimnach list "The Species of Epiphytic Cacti which was published in the Epiphyllum Society of America's Directory.)
The species in the ancestry of epiphytic cactus hybrids are nearly all tree dwelling (epiphytic), though few also grow on cliffs or rocks (saxicolous sak-sihk-uh-luhs). Unlike the better known terrestrial desert cacti, these epiphytes are generally native to forested areas, either perched in the trees with other epiphytes, such as orchids and bromeliads, or rooted in the ground and climbing up tree trunks, to which they adhere by means of aerial roots.
Epiphyllum: Haworth. 1812
About 12 species of scandent (skahn-duhnt meaning climbing)-pendent cacti distributed throughout most of Latin America. Stems flat or apically. Flowers usually nocturnal, rarely diurnal(staying open all day, as in E. crenatum and E. laui), funnelform, white to yellowish.
Aporocactus (a-por-oh-kak-tuhs): Lemaire. 1860
Two species of epiphytic or saxicolous ( sak-sihk-uh-luh), vining cacti with cylindrical, many ribbed, densely spiny stems. In cultivation, plants are usually grown in hanging containers so the stems are pendent. Flowers tubular, spiny, expanded apically, more or less unsymmetrical, petals red or purplish. Popular name: Rat-tail cacti". The genus is close to Heliocereus (hee-lee-oh-sehr-ee-uhs) but can be distinguished by its thinner, more ribbed stems and more or less zygomorphic flowers. (meaning-Bi-laterally symmetrical flowers, or flowers with only one plane of symmetry, ie. mirror-image right and left-hand halves, but with different top and bottom halves (assuming the flower lies horizontally))
Disocactus: Lindley. 1845 (Wittia, Wittiocactus, Chiapasia, Pseudorhipsalis, Bonifazia)
Twelve species of fat-stemmed spineless epiphytes native to many Latin American Countries. Flowers small, tubular to expanded, red to yellowish or white. Three groups can be recognized: section Disocactus with red, tubular to expanded flowers, and section Wittiocactus, with straight, short tubular, unexpanded flowers, the flowers of both sections being pollinated by hummingbirds, and section Pseudorhipsalis (soo-do-rip-sa-lis), with whitish, expanded, bee pollinated flowers.
Heliocereus (hee-lee-oh-sehr-ee-uhs): (Berger) Britt. and Rose. 1909
Four saxicolous or epiphytic, extremely variable species from Mexico, Guatemala, El Salvador, Honduras and Nicaragua. Stems flat to 3-4 angled, often spiny or hairy. Flowers funnelform, tube spiny or hairy, petals red, orange or purplish, rarely white. A large percentage of orchid cacti (Epiphyllums) have Heliocereus specious in their ancestry. The genus is hardly separable from Nopalxochia.
Hylocereus (high-loh-sihr-ee-uhs) (Berg.) Britt. and Rose. 1909
About 15 species widespread in Latin America. Stems clambering, triangular, more or less spiny. Fls.. usually large to very large, rarely small, nocturnal, the base usually spineless and with large overlapping scales, or rarely with small, separated scales and sometimes spiny (H. trigonus), petals usually white, rarely red (H. extensus, H. stenopterus).
Lymanbensonia: Kimn. 1984
A genus mainly differing from Rhipsalis by the presence of a floral tube over a centimeter long. The single species is terrestrial.
Nopalxochia(no-pal-ho-kee-a): Britt. & Rose. 1923 (Pseudonopalxochia, Lobeira)
A genus hardly separable from Heliocereus, from which it differs in its flattened stems and less spiny or hairy flowers.
Rhipsalis: Gaertner. 1788 (Acanthorhipsalis (uh-kan-thuh-rip-sal-us), Erythrorhipsalis (e-rith-ro-rip-sal-us), Hatiora, Lepisumium (lep-is-mee-um), Pfeiffera, Pseudozyggocactus (soo-do-zi-go-cactus), Rhipsalidopsis rihp-sal-ih-dohp-sihs))
About 75 species of pendent or creeping epiphytes from Mexico to southern South America, as well as from the tropics of the Old World. Stems flat to multi-ribbed, with or without spines or hairs. Flowers symmetrical (not zygomorphic), usually very small and yellowish-white,rarely to 2 cm long and reddish (R. rosea, R. gaertneri), fruits minute.
Schlumbergera: Lemaire. 1858 (Epiphyllanthus, Zygocactus)
Brazilian epiphytes with short jointed stems and long-tubed reddish, often zygomorphic (unsymmetrical) flowers.
Selenicereus (seh-lee-nuh-sih-ruhs): (Berg.) Britt. & Rose. 1909 (Cryptocereus (krihp-tuh-sih-ruhs), Deamia, Strophocactus (struh-fo-cactus))
A genus of some 10-20 species, widely distributed throughout Latin America from Texas to northern South America. Stems long and scandent, mostly many-ribbed (S. testudo, S. inermis, S. wittii) or flat and lobed (S chrysocardium, S. anthonyanus). Flowers small (S. innesii) to very large, nocturnal, funnel form, tube hairy, often spiny, petals white to yellowish.
Trichocereus (trihk-uh-sih-re-uhs) (Berger) Biccobono. 1909
A genus of some 40 species widely distributed in the Andes from Peru to Argentina. The stems are cylindrical and usually long, thick and spiny, while the large nocturnal flowers are white. All species are terrestrial except for T. arboicola, which is epiphytic in cloud forests. It is only distantly related to other epiphytic cacti. T. arboricola Kimm 1990. Bolivia. Stems eventually pendent, 2.5-4 cm thick, 9-11 ribbed, spiny. Flowers 12-13 cm long, opening 9-10 cm, petals white.
Weberocereus: Britt. & Rose. 1909 (Eccremocactus (ehk-ruh-moh-cactus), Werckleocereus)
A genus with very diverse stems--short and pendent or long and vining and flat to many-angled, but with similar, small, nocturnal, odd-smelling, bat-pollinated flowers. Some species are often included in Werckleocereus (3 angled stems) and Eccremocactus (flat-stems).
Care of Epiphyllums
SOIL: (From Rainbow Gardens)
A commercially packaged indoor plant mix, but it must be coarse and fast draining, this can be improved by adding some perlite, fine bark, pumice, or even small gravel. Avoid African Violet mix, has to much peat moss. For a home mix use: 1 part leaf mold, 1 part coarsely ground bark, 1 part 1\4"-1\2" redwood or fir bark, 1 part perlite (or horticultural pumice) you can also add 1\2 part horticultural charcoal. A commercially packaged camellia-azalea mix is a satisfactory substitute for leaf mold, as long as the mix is three parts organic matter and 1 part perlite or pumice. Rainbow Gardens does not use peat moss or vermiculite. For each cubic foot of mix add 1\2 cub bone meal.
Added note: In the "Cactus and Succulent Society of America's" Cactus and Succulent Journal Vol. 70 No. 1 January-February, 1998 there is a very extensive list of "Soil mixes for Epiphytic Cacti" on page 12
WATER: (From Rainbow Gardens)
Never let the mix of an established Epiphyllum dry out completely. Water thoroughly so that the water flows freely from the drain holes. Then allow the top 1\3 of the mix to dry before watering again. Clay pots dry out more quickly, plastic pots don't. Overwatering is more of a danger, so if you tend to overwater stick with a clay pot. Their natural growth cycle is in the spring and again in the fall and of course need more water at this time. Plants usually rest after flowering and again during the winter months. If the plant looks a little wilted from the stress of producing flowers do not increase the water. This is a natural phenomenon and the plants will usually recover their plump look by fall. In winter give just enough water to keep moist.
LIGHT: (Rainbow Gardens)
Filtered sunlight or a few hours of morning or afternoon sun, but never direct noonday sun. They may be grown under a lath, a fully leafed tree, or near a window where they will get morning or afternoon sun. Plants can be put outdoors until the cold weather arrives, when they are brought back into the house put in an evenly cool, but not freezing room where lights are not turned on after sundown. They can be grown under fluorescent grow-lights. The distance from plant to light should be at least 10 inches. The amount of light should be 500 to 1000 foot candles. The length of time to which the plants are exposed to artificial light should correspond with the actual hours of daylight during each season of the year. This is essential for bud development. Yellowish or sunburned growth indicates the light is too strong. Weak spindly growth with no evidence of a strong mid-rib indicates the plants are not getting enough light.
TEMPERATURE AND HUMIDITY (Rainbow Gardens)
They are very adaptable plants, but prefer temperatures between 45-70 degrees Fahrenheit. They will tolerate extreme heat if they are well shaded and the humidity is kept up. They will tolerate temps. less than 32 degrees Fahrenheit for a few hours, but will freeze if exposed to freezing temps. for any length of time. They are susceptible to cold damage when the temps. drop below 40 degrees F. In the house they like an evenly cool location 45-50 degrees F. with no artificial light after sundown. Cool temp. and long nights are essential for bud formation.
They like 50% humidity, but will tolerate less, a gentle misting is beneficial during the dry summer months to keep the humidity up. They like free air movement, but not gusting hot or cold winds.
FERTILIZING: (Rainbow Gardens)
They need a balanced fertilizer such as 6-6-6 or 8-8-8 or 10-10-10 at least once a month starting in spring and ending in fall. Do not use a fertilizer too high in nitrogen (over 10%).
In order to promote blooms and harden off tender young growth for winter, fertilizer with a low nitrogen or no nitrogen, such as 0-10-10 or 2-10-10. This is used once at the end of February and again in early November. Plants receive no fertilizer in Dec. and Jan.
PESTS: (Rainbow Gardens)
Scale and mealybug, control with insecticide soap or spray them with 1\2 rubbing alcohol and 1\2 water. Snails and slugs love epiphyllums and can do a lot of damage, snail bait should be put out on a regular basis.
BLOOMS: (Rainbow Gardens)
Takes 2-3 years to raise a blooming size plant from a rooted cutting. Day bloomers start blooming in late April, peaks in May and ends in June. Night bloomers are in late summer and early fall.
REPOTTING: (Rainbow Gardens)
Wait until about a month after blooming season to repot the plant, unless there is suspect rot or root pests. Loosen the root ball slightly and shake off some of the old mix, repot using fresh mix, don't water for one week after repotting, water lightly for a month or so then put the plant back on a regular schedule.
PROPAGATION: (South Bay Epiphyllum Society)
Dip cutting in a rooting compound, let the cutting set in a cool shaded place for a week or more so the cut end will callus. Plant the cutting an inch or more into the potting mix so that an areole on each side is below the soil line. It is from theses areoles that new growth will start. The potting mix should be loose with enough coarse material to keep it from compacting and provide good drainage. After the cutting is potted up, do not water, until roots have formed, it cannot take up water without roots and it will rot. Withhold water for a week or two then mist or sprinkle lightly. Begin to water gradually when you are sure that roots have formed, do not let it dry out completely. If the cutting starts to rot, cut off the rotted area and start over again.
Care of other Epiphytic Cacti: (Rainbow Gardens)
Culture, mix and blooming period of the "less spiny" epiphytic cacti, rhipsalis ( blooms on 1-2 year old plants), chiapora, cryptocereus, disocactus. nopalxochia is the same as for epiphyllums. The "spinier" epiphytic cacti Heliocereus, Hylocereus, Selenicereus bloom during the spring and on up into early summer on 2-3 year old plants. They prefer a well drained soil and a little stronger light than epiphyllums.
Aporocactus- Their care is generally the same as Epiphyllums, but the soil needs to be more pourus It can tolerate strong sunlight. It blooms in the spring,on 1-2 year old plants, or the same as epiphyllums.
Thanksgiving(by U.S. date) and Christmas cacti and Easter cacti
These thrive on a temp. of 50-70 degrees F. strong but indirect sunlight. They will do well outside in the summer hung in a tree but must be brought in before the temp. starts to drop in the fall. They will freeze if temps. drop below 32 degrees F. Thanksgiving and Christmas cacti bloom between these holidays and the Easter cacti blooms around Easter.
LIGHT: In habitat they grow at and altitude between 3000 and 5000 ft. they root in plant debris among the branches on decaying humus or on the ground in stony, shady places. They need bright but filtered light. If grown in the house keep away from a blast of a forced air furnace and away from the fireplace and cold drafts.
SOIL: A mix of 40% perlite and 50% peat moss (PH 5.0 to 6.0) is best.
WATER: In habitat, rainfall varies from 17 inches per month from Dec. to March, to 3 inches a month in the dry season. They require moisture at all times. Let them dry out slightly between watering, but never let them go completely dry.
FERTILIZER: Once a month with 10-10-5. Christmas or Thanksgiving stop fertilizing (September) 1-2 months before the short day period (or light span less than 12 hours). Start again after blooming is over. Easter cacti feed all year long except during blooming period.
INDUCING BLOOMS: For Christmas or Thanksgiving cacti only, Easter cacti do not require this kind of treatment. Temp. method: Keep nighttime temps. around 55-60 degrees F. Anything above or below will result in only growth. Photoperiod method: Short night period (less than 12 hours) at temp. above 60 degrees F. will initiate flower buds. This should take 3-4 weeks. Using either method, once flower buds appear, resume normal growing habits. Cooler or higher temp. during any stage of bud formation will delay flowering an additional two to four weeks.
Additional note from cacti_etc.: When new flower buds start to form, don't move the plant until the buds are near opening. If you move it when the buds are too small, they are likely to fall off.
Additional Info. Flowers that are white, yellow and pink will show color variations in their blooms if temp. drop below 50 degrees F. during any stage of the bud formation. Does not apply to Easter cacti.
Pests: Effected by snails or caterpillars. Do not use Diazinon, will cause stunting and distortion. For fungus rot use a good fungicide. Root mealy bug, submerge plant in Malathion until no more air bubbles are seen. Spotted branches or holes in branches is not uncommon, it is caused by uneven temps. Can only be prevented by growing in a perfectly controlled environment. It does not effect blooms.
Propagation: From stem cuttings may be done at anytime, but if done in winter it may require bottom heat 70 degrees F. Fert. after rooting has started 2-3 wks later. Pinch back in May or June to increase branching.
1. Cacti by Sir Oliver Leese, published by Triune Books, London, England 1973
2. Rainbow Gardens 1444 E. Taylor St. Vista, Ca. http://www.cactus-mall.com/rainbow_gardens/center.html
3. Jim Hunters' Epi Universe 'Night Blooming Cereus' http://www.epiuniverse.com/NightBloom.htm
4. Jim Hunters' Epi Universe 'Epiphytic Species http://www. epiuniverse.com/species.htm
5. Jim Hunters' Epi Universe 'What is an Epiphyllum? http://www.epiuniverse.com//whatis..htm
6. South Bay Epiphyllum Society, Palos Verdes, Ca. http://www.beachlist.com/epif/epi2.html
7. 'The Species of Epiphytic Cacti' by Myron Kimnach
8. 'The National Gardening Association Dictionary of Horticulture', Viking Press, Produced by The Philip Lief Group, Inc.
9. 'Hamlyn A-Z of Plant Names' by Allen J. Coombes, Chancellor Press, 1994
10. 'The Encyclopedia of Cacti' by Dr. Willy Cullmann, Dr. Erich Gotz, and Dr. Gerhard Groner, Timber Press 1987.
From the San Francisco Cactus and Succulent newsletter
Fat Plants of South America
By Carl Frederick (San Anselmo, Ca.)
Of our Southern Hemisphere, South Africa is brimming over with succulents of all kinds, Madagascar appears to have a large number of species with many more undoubtedly yet to be discovered, yet Australia, mysteriously, has few. South America appears to have fallen in the middle with regard to numbers of species, boasting many types of cacti and a much smaller number of non-cactus succulents. Of these, there are a number of succulent trees that are not well known, even in their native countries.
Recently I have been trying to find out more about these species since I seem to have picked up a bug for succulent bonsai and the infection appears to be permanent. Admittedly, I haven't done a major search through the literature but a perusal of our wonderful nurseries here in the states, as well as those of Europe and South Africa has left me with the impression that virtually none of these plants are in cultivation. The one best known and available is a variable species, Chorisia speciosa, which appears in Jacobsen's Lexicon of Succulent Plants. In fact, probably the most famous picture of him that appears in his Lexicon as well as in Rowley's Caudiciform and Pachycaul Succulents, shows him leaning up against a C. speciosa. The most striking thing about this plant is its thick covering of stiff, conical spines. The tree is quick-growing but is possible to bonsai as evidenced by beautiful plants turning up at shows. Chorisia also contains C. insignis that I have never seen but from the description in nursery catalogs, is similar to C. speciosa (perhaps it's synonymous?). Another species, probably not in cultivation, C. glaziovii, has a more succulent, bottle-shaped trunk. The genus Ceiba also offers succulent species including C. pubiflora with a bottle-shaped, thorny trunk and C. jasminidora with thick roots and a squat trunk up to 2 meters in diameter. The giant of the clan is Cavanillesia arborea which has a Baobab-like habit.
When the incredibly prolific ethnobotanist Richard Evans Schultes was searching for a good source of rubber-bearing trees in South America during WWII he explored some of the fascinating mountains rising up out of the jungles of the Amazon. Some of these mountain tops represent very different ecosystems than the jungle floor below, often consisting of grassy, xeric mesas bordered by precipitous cliffs. While in Colombia, Schultes had an opportunity to botanize a range of these mountains called the Sierra de Chiribiquete. Upon one of them, Cerro
Pictured: Cavanillesia arborea , in habitat, Caatinga, Brazil Chiribiquete, he found stunted plants with glabrous, leathery leaves coated with thick waxes or a dense pubescence. There were epiphytes with hugely exaggerated pseudobulbs and squat, fleshy rosettes hugging the ground. Two of the plants he named suggest the impression that they had upon him: Graffenrieda fantastica and Vellozia phantasmagorica. He even found miniature rubber trees, later called Hevea nitida var. toxicondendroides. These rubber shrubs were, unfortunately not of any economic value as sources of latex but I have the feeling that some of us would still find them and the other xeric flora he encountered...valuable
Let's save our pennies, get some permits, visit these places, and collect some seeds. There's still a world of unknown succulents out there waiting to be appreciated.
Wade Davis, "One River", 1997.
Marlon Machado, private communication, 1999.
Grigsby's catalog, 1999.
Cacti of Brazil
By Tom Steppe (Alameda, Ca.)
Cacti of Brazil: Melocactus, Discocactus, Uebelmannia, Pilosocereus and Gymnocalycium. Melocactus is an old name, one of the ten or so brought back to Europe by the early explorers and made known to Linneas. They are usually globular, solitary plants, with prominent ribs and spines. Many small flowers are usually produced each year on mature plants of the cephalium, a specialized flower-producing area on the top of the plant. The Genus ranges from the West Indies, Mexico and Venezuela all the way to Peru and tropical Brazil. M. intortus grows in the US Virgin Islands. The largest concentration of species are in the Brazilian states of Bahia and Minas Gerais. They are tropical growers, suffering below 60 F.
Discocactus are more recent. A few species were known to Britton and Rose, but most have been discovered since the 1950's. They are also globular, usually solitary, developing the prominent cephalium of the 'Melo' with time and maturity. A few species are from Bolivia and Paraguay. Most occur in Bahia, Goias and Minas Gerais and border areas. They also suffer below 60 F. The genus Uebelmannia is named after Uebelmann, the Swiss grower. They are known well only since the 50's. Most have ribs and spines resembling a Notocactus, but are extremely-shy flowering plants and much more difficult to keep alive on their own roots. All are from Minas Gerais and Bahia and border areas. They suffer badly below 60 F.
Pilosocereus is a tall growing genus, with species from 3 ft tall to more than 20 ft tall. They are prominently ribbed and tall growing, hence the cereus in the name. A few sport a pseudocephalium, or side-growing cephalium. The genus Pilosocereus ranges from as far north as Barbados and the Windward Islands, but most species are from Bahia and Minas Gerais in tropical Brazil. They are wide-ranging, but 55 F or higher should be used for best growth. The genus Gymnocalycium is a broad ranging one, mostly from Argentina, Paraguay and Uruguay. A few species occur as far south as Patagonia and a few occur into southern Brazil. Those Brazilian Gymno's include G. denudatum, buenekeri, horstii, netrelianum and uruguanense. A few well-known grafts are in the trade as 'Hibotan' and 'Nishiki', the grafts of variegated G. mihanovichii. I'm sure we've all seen 'Red Cap', 'Pink Cap' & 'Yellow Cap' in the supermarkket, grafted on Hylocereus stock. The Gymno's are popular because they are easy to grow and easy to flower with most collectors. Some will even tolerate a light frost if kept dry.
Cacti, Clives Innes and Charles Glass
Gymnocalycium, a Collector's Guide, John Pilbeam
Growing the Semps
by Lucille Tobiassen (San Francisco, Ca.)
Sempervivums are easy alpine plants to grow. It is not fussy about the soil mix as long as there is good drainage. Common names are 'he & chicks' or 'house leeks' but there is no smell of onions! Sempervivum means "live forever" but many of the plants are monocarpic-meaning that the rosette dies after flowering and producing one crop of fruit. The flowers remain as the off-sets form at their base making it seem like the plant is immortal. Sempervivums have been in cultivation for a long time. In Europe they were planted on roof tops and beleved to keep demons away from the home. It was also thought they would protect the house from being struck by lightening. Horticulturally speaking, there are two distinct groups of Semps: tender species from Northern Africa or Canary Islands and the hardy species from the mountains of interior Europe. Many of the fancy hybrids grown today are derived from the European natives.
Botanically speaking, there are four plus types of semps. (1) tectorums are smooth and slightly hairy; the cultivar 'Ohio Burgundy' has a large rosette and develops a deep rich burgundy color. (2) arachnoi-deums are the "cob-web house leeks" and can tolerate a lot of sunlight. (3) calcareum are velvety and prolific, apple-green colored. Examples are sp. bicolor and sp. rubrum. (4) ciliosum are the tight rosette plants with many fine hairs-very exquisite. Jovibarbas look very much like semps but have yellow billed shaped flowers. The mother plant sends off the new plants on stolons radiating from the base. The plants sometimes break off the stolons and roll away to establish in a different location.
Semps are best grown with filtered light or in partial shade. However, many can tolerate bright sunlight once they have established into there growing medium. Propagation is simple because offsets are produced at the base of a stolon and usually develop roots readily. They can be removed and planted right away. This is one group of plants that you generally don't wish to see bloom for that signals the mother rosette to send up off-sets before dying. Small off-sets at the base of a plant should not be disturbed until they reach an ample size and the mother rosette has nearly dried up. Sometimes semps will bloom and die without sending up off-sets.
Martin and I grow over 250 different varieties of semps and we try to keep two heads of each to keep the propagation going. We have not distributed any Jovibarbas yet but plan to offer them for the big Strybing sale in May. I have successfully divided a Jovibarba luckhoffii 'Beacon Hill.' I also cored a Jovibarba 'Jungle Fires' that produced off-sets.
It all started from a generous gift of 30 flats from Bob Murray who raised many from seed and off-sets. I have been doing a little research on these plants and two years of propagation and sales have added hundreds of dollars to the plant sales. Finally, pests are rarely a big problem but root mealies are the most pervasive. Sometimes ants love to make a nest under a carpet of these plants. Enjoy your semps!!
San Gabriel Valley Cactus and Succulent Society
Succulent of the Month January 2000 - Gasteria
by Tom Glavich
Gasteria is a popular and easily grown genus of plants that has been collected and kept as house plants since the 17th century. Gasteria are readily distinguished from Aloe and Haworthia (both very close relatives) by the shape of their flowers. Gasteria flowers are carried on a long scape (leafless flower stalk). They generally hang down as they open, and have a swollen (gasteriform) portion at the base of the flower. The word Gasteria comes from the Greek "gaster" meaning stomach or belly.
Gasteria vary in size from about an inch to several feet in diameter. Almost all plants start out with the leaves in two rows (distichous), although most eventually begin to spiral, either remaining in a distichous spiral, or changing into a rosette form. Gasteria are very variable in appearance. They have definite juvenile and adult forms, have local variations in appearance, and have form and growth habits that are dependent on the soil type and amount of sunlight.
Typical natural habitats are humus rich, sandy soils. They grow on dry rocky hillsides and generally grow under larger shrubs, particularly when young. They also can be found in rock fissures or in the shade of large rocks. The roots are shallow and thick, and this helps the plants obtain moisture from barely wet soils, and nourishment from decaying leaves and debris from larger shrubs.
Gasteria cultivation is easy. They are mostly winter and spring growers, but exhibit some growth all year except for the hottest part of the summer. They are tolerant of almost any growing mix, although they do best with high organic content mixes, similar to their natural growing conditions. They prefer partial shade, particularly in the afternoon, however the best color is obtained by giving them as much light and sun, short of sunburn, as possible.
Gasteria are generally free from most pests. The one difficulty is 'black spot', a fungus that attacks many Gasteria, particularly large, show quality plants. The fungus is rarely fatal, but causes large unsightly black spots on the leaves. There is no way to remove the spots, and since the leaves remain on the plants for several years, the fungus can quickly ruin show plants. The fungus can be minimized by keeping the leaves dry, and particularly keeping dew off the leaves. It can be prevented by regular application of systemic fungicides, but vigilance is required. Funginex is a popular rose fungicide that works well on controlling black spot.
Gasterias are readily propagated from offsets at the base, which can be simply pulled off and planted. Leaf cuttings will also root easily. Gasterias left in the open in California will be rapidly pollinated by hummingbirds, although the resulting seed is then an uncontrolled and unknown hybrid. Controlled pollination is easily accomplished with a small brush, or toothpick Seed can be collected as soon as the fruits start to dry.
Gasteria hybridize easily, and cross pollination can produce interesting plants. A number of hybrids are available, as are several variegated and at least one monstrous cultivars. Intergeneric hybrids with Aloes and Haworthias are also available.
Selected species include:
Gasteria nitida var. armstrongii (usually found as Gasteria armstrongii) This is a small plant, with distichous leaves only a few inches long. The leaves are dark green (almost black in some cases), and are tuberculate (covered with small bumps) It offsets freely from the base, forming nice clumps. Larger specimens can be obtained by over-potting.
Gasteria batesiana is another of the dark green tuberculate species. The leaves spiral with age to form rosettes. The plant is much larger than Gasteria armstrongii, and the tubercles are often a lighter green than the main body, giving the plant a speckled appearance.
Gasteria carinata var. verrucosa (often found as G. verrucosa) is a lighter colored species. It is larger than G. batesiana, with thinner leaves, The tubercles are much larger, and often merge into large areas of glaucus green on a dark green background.
Gasteria carinata var. carinata cultivar 'Grat Brak' often found as G. schweickerdtiana is a particularly beautiful plant. In this cultivar the tubercles on the underside of the leaves form bands. Tubercles also merge on both the upper and lower leaves to form glaucus green stripes on a dark green background.
E. J. van Jaarsveld, Gasterias of South Africa
F. Sajeva and M. Costanzo, Succulents, The Illustrated Dictionary
Gates Cactus And Succulent Society
By Dennis Kucera
There are about 60 species of Sansevierias belonging to the Agave family. Before the 1970's they were included in the Lily family as found in the older Exotica plant encyclopedia. They have clusters of small fragrant white flowers with 6 lobes like a lily. Most species are stemless, xerophytic succulents, with rhizomes, underground stems as big as your finger that end in a cluster of upright leaves. These evergreen perennials grow wild in dry rocky habitats in tropical and subtropical Africa, Madagascar, India and Indonesia. They are frost tender and should be grown in a protected shaded patio, warm greenhouse or as a house plant. Many kinds can tolerate neglect and only need ccasional watering once a week or less in summer with good drainage. A half strength liquid fertilizer can be given monthly during the warmer months. Propagation is easily done by separating rhizomes with leaf clusters and also by leaf cuttings.
S. trifasciata, the mother in law's tongue or snake plant and its varieties is the most commonly grown. It is native to Transvaal, Natal and the eastern cape of South Africa. This species has lance shaped, fleshy leaves to four feet high with horizontal light and dark green bands. S. trifasciata 'Laurentii' the most popular variety has upright leaves to 18 inches with golden yellow marginal stripes. The 'Golden Hahnii' variety forms dwarf rosettes of broad leaves to 8 inches long, with wide golden yellow vertical stripes. The variety 'Silver Hahnii' is a birds nest type with leaves with metallic pale silvery green leaves. S. Kirkii of horticulture is probably S.grandicuspis from the Congo. It has coved leaves about one inch wide to 20 inches long of deep green, cross banded with light grayish green. S. singularis has cylindrical gray brown or reddish leaves to 25 inches or more. It comes from tropical Africa. S. gracilis has stoloniferous stems sprawling above the ground with clusters of narrow pointed leaves to 20 inches long. The leaves are cross banded light and dark green. It is native to eastern Africa. S. deserti is called rhino-grass and grows wild in the Kalahari desert to the dry veld, Transvaal and Rhodesia. It forms dense masses of very stout cylindrical leaves 1 inch thick and 3 feet long with a very sharp tip. It is avoided by wild animals.
Kaktos Komments - Houston
Mammillaria bocasana - "Fred" by David Thomas
Name: Mammillaria bocasana - "Fred"
DISCUSSION: The cactus named Fred has been around for more than 25 years. For those who think Fred is a frivolous name for a cactus, know that nurseries such as Grigsby's Cactus Gardens list Fred in their catalog between the Fraileas and the Gymnocalyciums. Various stories have circulated on what genus this plant belongs to, with claims that it is either an Echinocereus, a Cophophora, or a Mammillaria. One prominent Mammillaria authority threw Fred out of a Mammillaria symposium claiming that the plant was a Crassula.
A non-specialist nurseryman grafted his stock Freds in order to produce large numbers of plants but got just the opposite reaction. The rate of growth slowed, the plants tightened up and, out of 100 grafted plants, two produced normal heads of Mammillaria bocasana complete with typical hooked spines, wool and flowers.
It is assumed that Fred does not occur in habitat, but was a monstrous seedling or developed as an offset at some time in the past. There is no record of who the original grower or propagator was.
DESCRIPTION: A monstrous cactus that sometimes will have crested sections. The straight, weak spines, when they do appear, are about 5 mm long and range from 3 to 6 per areole. Flowers are rarely produced and usually abort before opening. Those flowers that do open are small, cream colored and very Mammillaria like.
SOIL: Very loose mix.
WATER: When plant feels flaccid to the touch.
LIGHT: Strong to medium light. Direct light will turn the plant red.
REFERENCE: The Cactus File - British Journal
THE NORTHERN CACTOPHILE by JIM ALGER Saturday morning has come, I climb the stairs and water Time for my weekly duty, The ones in the hall window, Watering my cactus friends Then into the east bedroom -- So that they will smile at me Ones that dominate those sills. another week. I paddle down in slippers Finally I descend to And pour a dash of plant food The basement, a west window, Into my watering can Water the big Opuntia box, And head down to the basement. They're tough, for a less nice place. I put the long-spouted can I fill the water can Into the old tin wash tub For the next big watering (To keep a minimum mess), To let it breathe out gases, Fill it from the rain bucket. Warm to room temperature. The last rain water this year. The week's watering is done. Upstairs again I water Actually, I must admit The long cactus plant boxes Last weekend was so busy That dominate the dining room -- With visiting guests and all Yes, it takes real patient folk I missed their good watering, To live with a cactophile -- So it has been two weeks now, Neatly transplanted inside But they are so forgiving, From my summer rock garden. These friends from the great deserts. Pleasant smell of watered earth Now let the winter winds blow. Comes to me as I'm pouring. They are safe inside with me I stand back and view them all, Until the spring transplanting Healthy from this summer's heat. Into the garden again.
How Cold Is Cold?
Roy "Frosty" Paramore
One of the most irritating things about succulent literature is that the authors seldom agree with each other on how to grow plants. I can understand that differences in habitat in England, California and Texas make a "one for all" soil mixture impossible. But temperature? Should this be the same for every plant?
Miles Anderson, in his latest book CACTI & SUCCULENTS, gives more information on minimum temperatures for cacti and succulents than I have ever read. I have listed a few of the more popular genera and species:
Ancistrocactus ... down to 10 degrees F. for short periods
Ariocarpus ... down to 15 degrees, if dry
Most Astrophytum ... 20 degrees, briefly
Coryphantha ... 10 to 25 degrees, briefly
Most Echinocereus ... 15 to 20 degrees, briefly
Epithelantha ... to +/-20 degrees for brief periods
Mammillaria ... usually not less than 20 degrees, dry
There are many others, and this book is in our library. I have often wondered who keeps the Chihuahuan Desert plants warm in winter.
The Cactus Wren
The cactus wren became the Arizona state bird in 1931. It is the largest member of the wren family but is very different from its relatives. The cactus wren has a brown cap; heavily spotted throat and breast; a white line over its eye; and white spots in the outer tail feathers. Even it's song is different. It is a monotonous chut-chut-chut.
Nest building is one of the cactus wren's most interesting activities. A great portion of each year is spent either building or replacing a shelter nest or a brooding nest. The basic structure of both nests is an eight to ten inch long tunnel with a ball at the end made from straw-like plants. Although the cactus wren prefers to build these nests in the tops of cholla and prickly pear plants, the nests may also be found in Joshua trees, yuccas, saguaros, mesquite trees, ironwood trees, and Palo Verde trees.
The cactus wren does not migrate for the winter. To shelter itself from the cold, the wren will build the basic nest structure and line it with whatever is locally available to make the nest warm and padded. Nests have been found lined with chicken, quail, and dove feathers; sheep wool gathered from mesquite thorns or barbed wire; cotton; paper towels; and pieces of clothing.
The spring brooding nest usually has thinner walls and less lining than the shelter nest. Wrens will often build three or four extra nests close to a brood nest. These dummy nests confuse the cactus wrens' enemies such as rattlesnakes. They also shelter the adult bird that is not brooding the eggs and the young birds old enough to leave the brooding nest.
Although building nests in cacti provides the cactus wrens with some protection from their enemies, the cactus spines are a hazard to the birds. Generally, the birds come and go with ease but every year some get caught by the spines and perish. Usually these are young, inexperienced birds but occasionally an adult will be found entangled in cactus spines.
Cactus wrens have adapted to the changing environment in Arizona. Today, they are found in the cities as well as the cactus-covered countryside.
Arizona Highways; March, 1948 (yes, an old magazine)
Spinal Column - Michigan
by Lou Kilbert
The Mesembryanthemaceae is a large family of plants confined, almost exclusively, to South Africa. These plants range from dwarf shrubs to tiny plants that some would need a magnifying glass to see. They are mostly "leaf" succulents; that is to say that they have special cells designed to store water inside the leaves. A few have underground, carrot-like, water storage tap roots that may be raised above ground in cultivation for display purposes as caudiciformes.
The larger growing species make excellent landscape plants in California and many highways are lined with displays of their brilliant daisy-like flowers. We find it difficult to flower the larger shrubbies, but many of the tiny shrubs make excellent pot plants.
The most popular genus is the Lithops or South African Living Stones. There are approximately 80 species listed in Jacobsen and over 445 varieties listed by Steve Brack of Mesa Gardens listed in his seed catalogue!! (Jacobsen devotes all 525 pages of volume 3 of his Handbook of Succulent Plants to the family; that book is in our library.) Although everyone seems to love them, many find them difficult to grow. Watering can be tricky. They need water in spring and fall but should be kept dry in the winter and, depending on the situation, during the summer as well. They flower during the fall. The flowers are usually about twice the size of the plant body, which consists of one pair of leaves. The larger the pair of leaves, the larger the flower. They are either yellow or white and never pink, red or blue. The petals are very thin giving an informal appearance to the flower. The flowers open at about the same time of late afternoon. This diurnal rhythm is quite fascinating. I noticed that the flowers opened at the same time even when under lights in the center of an enclosed shopping mall where they were hundreds of feet from the nearest glimpse of daylight! Many mesembs have scented flowers, but I have never detected a fragrance from Lithops flowers. However, Haage states that L. lericheana (a variety of L. karasmontana) has "sweetly fragrant flowers".
When you press a finger to your Lithop, it should feel like a piece of rubber. It won't feel as hard as a rock; but it definitely should not feel soft and squishy, which would indicate overwatering. If you've lost them to overwatering before, wait until the skin starts shriveling; that may take several months! South Africa is a "land of light". The light is very intense. If you can't supply enough light in winter, use fluorescent lighting with the plants almost in touch with the tubes.
For those experienced in growing plants from seed, mesembs are relatively easy and one year old seedlings can be easy to handle. The seed is very tiny and it's always surprising how large a seedling can jump from such humble beginnings.
Other genera have very different habits. Many are summer growers. Some only grow for one or two months each year and "rest" for the remainder. Learn as much as you can about your plant so that you can give it what it wants when it wants it. They all need good quality, well draining soil and food! I've seen many people starve their plants, thinking somehow that "desert" means "starvation". That only produces weak plants that are short lived and don't flower. Why don't you come to the meeting and learn more about these fascinating plants.
by Lou Kilbert
Notocactus is an interesting genus of small barrel cacti for the beginner. Perhaps, the number one problem for beginners is overwatering. Surprise, Notocacti will suffer if they do not get some water in winter. If Notocacti shrivel, they take a very long time to recover and are unlikely to flower during that summer. Summer flowering, that's another reason Notocacti may be a good bet for beginners. Their flower buds are formed in spring and bloom in summer, unlike many cacti that require either cold temperatures or long dark nights in winter to bloom in spring. If you have difficulty giving adequate light in winter, Notocacti can still flower if you put them out-of-doors starting about late April. Outside, part-shade is probably best for Notocacti.
Adding a little extra organic matter to the soil is a good idea when potting these cacti. I've grown them successfully in both plastic and clay pots. They need adequate fertilizer to do well. The majority fall into the 2-3 inch range; although, some can get to be much larger, to three feet tall. Most produce flowers in various shades of yellow, but some are pink, purple and even red. There are so many petals that some flowers look "double". Many of these flowers have that glossy sheen that makes them glow "like waterlilies". Each flower bears a "beauty mark" in the center. The stigma (female part) is red from brilliant to a dark and velvety burgundy.
They are easy to grow from fairly large seed that are easy to handle. It will take about three years for the small varieties to bloom from seed and at least 5 years for those that are larger growers.
Some notable and popular species: N. ottonis blooms quickly from seed (2 years?) and can start when it's only one inch in diameter. It forms a nicely shaped dark green globe with harmless, but beautiful, spines that may be bright yellow to brilliant red in color. My favorite variety has soft, long twisty spines that are red in color. I can grow many of these in my collection because there is such a great variety in spination. There is at least one variety with reddish flowers (N. o. Venclusianus). I would give this five stars out of five as a "best cactus for beginners". N. uebelmannianus looks a lot like N. ottonis but has a wooly tuft of white fur in the center. It's notable for the beautiful purple flowers that are fairly freely produced. The species that fit into the subgenus, that used to be given full status as the genus Malacocarpus, also produce a lovely tuft of white fur in the center of a plant with many dark green ribs.
N. scopa is another plant that I can grow in quantity because of its variability. In its best clothes, it is a white furry ball with glowing ruby red spines sticking out like Christmas tree ornaments. It may not flower freely and the flowers are relatively small for the genus. I have selected a variety from seed that clusters with heads about 1/2 inch in diameter; each little head is covered with dense white wool.
N. lenginghausii may get to be one foot tall in a pot (to three feet in habitat). It has brilliant yellow spines and the top of the plant is very unusual, it always faces towards the sun. The plant grows as an upright column with the top like a face placed almost vertically towards the sun.
Other species worth growing: N. mamulosus blooms freely, N. submamulosus has strong sharp dagger spines, N. buiningii has bright blue skin studded with brilliant yellow spines sometimes highlighted with dark red. There are many other species, all excellent for the beginner.
St. Louis - Henry Shaw C & S
by Pam Schnebelen
I started growing Haworthias almost 20 years ago. At that time, I lived in a city apartment with no yard, few windows and no space for large plants. Haworthias did well in my limited environment. They are easy to grow.
Haworthias are small and compact, most fit comfortably in three to five inch pots. They grow as solitary or clumping rosettes that stay close to the soil level. They don't have to be outdoors in summer and they don't need a hard, cold, winter dormancy. Instead, most Haworthias are winter growers. They are tolerant of soil mixes and watering schedules. And, they bloom easily, putting up cute whitish flowers on long stems that tower above the body of the plant.
My first lessons in propagating from offsets and leaf cuttings came from working with Haworthias. On top of all these growing characteristics that make these plants ideal for both the beginning and advanced collector, Haworthias are absolutely gorgeous. They have fascinating shapes and symmetries, ranging from tight spirals of layered triangles to heaping mounds of tiny clear-skinned grapes. Colors vary from bight yellow to a deep red-brown, from pale lime green to deep blue green.
Leaf textures are interesting too. Some plants have bright white raised tubercles in stripes or patterns of dots on the tops and bottoms of leaves. Other leaves have rough surfaces that look as though they have been dusted with sugar crystals.
Then there are those "windows", perhaps the most interesting leaf feature of these plants. Many Haworthias have clear leaf tips and surfaces. These translucent windows allow sunlight to penetrate into the body of the leaf. In most plants, photosynthesis occurs only on the surface; in windowed plants, photosynthesis can occur throughout the body of the leaf.
When discussing odd plant features and Haworthias, we must also discuss roots. Haworthias have long, fleshy, "contractile" roots. In the cool wet seasons, these roots serve as a water store, much like the caudiciforms. In the dry hot summers, these roots serve two functions: they return moisture to the body of the plant and, while doing so, they shrink, pulling the plant deeper into the soil to protect the plant from sun and heat.
by Jonathan Watt
Copiapoa. A member of the cactus tribe with a very narrow natural distribution, occurring in the coastal region of Chile. The genus is named after the province there, Copiapo. There are 14 described species. The plants are simple globose to elongated cylinder in shape, depending on the species. One species is clumping, with many offsets. All varieties have prominent ribs, some have additional "woolly" growth on top of the plant, this is where flowers arise. The flower colors are yellows, with some tinged red and are funnel shaped.
What attracted me to this genus is the apt description in the catalogs; light green colored, blue green colored, chalk white colored with black or dark spines!! What a combination! These are truly stunning plants to add to your collection. Copiapoa is a true desert plant. They are supposedly difficult to flower in cultivation because of the light requirement. So what?? Drainage in the potting mix is the most important factor, especially in winter. They need to be absolutely dry in winter and cooler temperatures, but still in full light. They can be watered freely in summer, allowing a dry out in between, but watch it in cooler months.
Here are some to try: C. coquimbana; a clumper with black spines, wool and yellow flowers. C. echinoides, an elongate cylindrical shape, blue/green color, black spines, woolly at the top, yellow flowers. Yow, does it get any better? C. cinerea, same as above in shape but chalk white in color with glossy black spines. A thing of beauty.
Give these plants a try so you can come up with your own superlatives!
by Jonathan Watt
Turbinicarpus: the word comes from Latin, meaning, "whipping, top-shaped" referring to the shape of the plant. There are between five and ten species recognized, all native to Mexico, especially the Chihuahan desert. They are relatively new to science, having been described within the last 50 years. But here's the rub, at various times Turbinicarpus has been included in Strombocactus and Pelecyphora and some species are still being moved around.
The plants are quite small and grow slowly. They are globe/cylindric in shape and generally quite spiny, except for T. schwartzii and T. schmiedickeanus, which look like Lophophora cacti.
Turbinicarpus flowers range from white, pink, lavender and yellow depending on variety. Some are striped with two colors. All flowers arise from the top of the plant out of the areole and flower in late winter/early spring, a good time for cactus flowers.
These plants are known for their tap roots which make them tricky to pot and keep alive. They require a very minerally soil, very good drainage and very careful watering in summer, none in winter.
Grow them in bright light, no direct sun, and a decidedly cool winter temperature of 30-40F is fine.
Here are some varieties to try: T. klinkeranus, white flowers. T. machrochele, white flowers with pink. T. pseudomachrochele, white flowers with pink. T. schmiedickeanus, rose colored flowers. T. kranziana, white flowers. T. schwartzii, white/lavender flowers.
The German botanists have been very active in researching these plants, hence the Teutonic species names.
by Jonathan Watt
Cephalocereus senilis: this month's plant of the month from the cactus tribe is one of the most recognizable cacti to anyone, grower or not. The common name is Old Man Cactus, owing to the long, white, shaggy hairs completely covering the plant. The plant is the sole representative of its genus--a monotype.
Old Man Cactus is a columnar growing cactus, rarely branching. In nature, it can grow to 40 feet tall but in collections rarely to 10 feet. It flowers seldom in cultivation, forming a true cephalium that starts out on one side of the plant and eventually encircles the stem; having longer, more bristly "wool" and a creamy color.
Old Man Cactus is a slow grower and grows well on its own roots, hence they are mostly grown from seed. They can be grafted onto a suitable rootstock to speed up growth. The plants require a sunny location and coarse soil mix to thrive, with gypsum added to keep it on the alkaline side. A dry winter rest is essential, as is warmth and sun.
Try this Mexican native if you want an instant conversation piece for all your non-plant friends.
by Jonathan Watt
Agaves are members of the Agavaceae family. They are closely allied to the Liliaceae (Lily) family and were even classed in the Amaryllicaceae family until the 30's and 40's when they were given their own.
These plants are well known for their fleshy, wickedly pointed leaves (wear goggles when working with them!) and beautiful, much anticipated flowers. Some are true "century" plants, blooming only once during that time, then dying. Other species are more accommodating, flowering sooner, but alas, they ALL die after they bloom. some send up new shoots from the roots, some have bulbils that fall to the ground from the fertilized flowers and of course, there are seeds.
There are many species to try. Most can fit into an average collection though some are huge and require careful siting because of the spines and the high light needs. Agaves are very water sensitive. Do not over water, especially in winter. They do not seem to be picky in regard to soil.
Some to try: A. victoria-reginae, white edged leaves, stunning! A. triangularis, prominent ragged spines on leaf edges and tip, dark-green, wow! A. parrasana, a beautiful blue one with black spines, ultra cool! A. geminiflora, interesting threads on leaf edges--outasite!
All of these plants mentioned are native to Mexico. Many Agave are hardy into extensive regions of the U.S. It would be interesting to try some here in St. Louis. Those of you who have traveled to warmer places know the giants of the family. A. americana can be five to six feet tall and wide and then have 20 foot flower stalks. What a plant!
The Cactus Patch
Travels With Rob
November is a great time to visit Mexico and catch the Ariocarpus in bloom. This last year I traveled there on a plant photography expedition with Mike Navolio and two other friends, Larry Nicholes and Jim Boehmke. We met in Laredo Texas, where we crossed into Mexico, and headed towards the Sierra Madre Oriental and the Chihuahua Desert. We traveled in two cars; always good insurance against possible breakdowns and flat tires. We brought along all our food, camping gear, and detailed maps marked with locations of cacti that had been gleaned from friends and back issues of the Cactus and Succulent Journal.
The most frustrating part of any trip to Mexico is getting through the confusion and lines of the Mexican customs bureaucracy, changing money, and taking care of last minute details like getting ice and car insurance. But by midmorning we were speeding south towards Monterey on one of the fast, new (and expensive!), toll roads that are making travel there so much easier than just a few years ago. In much less time than I could have believed, we were past Monterey, past Saltillo, and climbing the grade to the high plateau of central Mexico. It was there that we left the highways, and took small, often unpaved roads into the remotest parts of rural Mexico.
Wherever you look in Mexico there are interesting succulents and cacti. Every stop is different and exciting, but for me, the real thrill of the trip was finding both Ariocarpus bravoanus v. bravoanus, and v. hintoni. These two plants are so new that they are virtually unknown in the US. But just as good was Turbinicarpus alonsoi, which took us into entirely new terrain for me - misty pine mountains, and deep, deep gorges, where we also saw Mammillaria schiedeana and Bombax ellipticum. We found several other interesting Turbinicarpus sites, many Ariocarpus retusus locations, and an A. agavoides hill that was new to me (we were helped with that one by a shepherd - it helps to have a Spanish speaker along). The agavoides were flowering, as were the A. scapharostrus. And we scouted out an amazing location for A. trigonus containing hundreds of plants. We never did find the limits of that population. We also saw Obregonia, many different Mamms, Feros, Corys, Thelos, Gymnos, Astros, and a Neolloydia. Thinking back on it, its hard to believe that we could see all that in just over a week. Oops, forgot to mention Calabanus! Not to mention Aztekium ritteri, and various Burseras, Cycads, Yuccas, and Agaves. Mexico is just that way- incredible.
We saw some of the destruction caused by illegal collecting. (ALL cacti collecting in Mexico is illegal.) On an isolated hilltop, after a long climb, we were only able to find a few plants of Turbinicarpus panarotoi. But evidence of collecting abounded, and it was plain to see how the greed of commercial collectors had nearly stripped this hill of its recently discovered new species. Populations of other plants seemed to be having better luck. I had been concerned especially about the Ariocarpus bravoanus v. bravoanus, which I had heard had come under heavy collecting pressure. But the colony we found seemed intact, as did that of v. hintonii. There also seemed to be a shift in the attitudes of some of the Mexicans we met. Whereas at times in the past I had been met by villagers attempting to sell me plants from gunny sacks they had filled, at several locations on this trip the locals were protective of the plants, and suspicious of our intentions. This is a good development.
The trip was wonderful. The weather was good, we had no hassles with the authorities, and we were so successful in finding the plants. It was a first for Mike, but I'm sure, not the last. And already, I'm starting to put together a mental list of plants and destinations for next time.
Cactus Courier - Palomar C & S
How Cactus Made It To The Americas
by Stan Yalof
Everything comes from somewhere. The Cactaceae are a New World only family, but where did they come from and how did they get there? We do know that Cacti, together with Didieras, Mesembs, Portulacaria, Ceraria, and Anacampseros, are closely related within the taxonomic order Caryophyllales. Didiera is found in Madagascar; Mesembs in south and east Africa; Portulacarias and Anacampseros in southern Africa. Tracing back from their present locations, I would place the ancestral home of this order in the vicinity of Mozambique. In this article, I have attempted to trace the movements of the ancestors of Cactaceae from Pangaea to South America and into North America. I have drawn from evidence in several fields, from the works of others, and from my own studies in piecing this story together.
During the Paleozoic Era, between 550 and 250 million years ago (MYA), most of the Earth's land masses had combined into a supercontinent which we call Pangaea. In the early days of its assembly, back in the Devonian Period some 500 MYA, the climate of the then smaller Pangaea was modulated by the influence of the ocean. Into this world life first stepped onto land. Beginning with a green dust, algae, which surrounded wetlands and the border of lakes, Bryophytes (mosses and liverworts, plants without roots), arachnids, and fish-like amphibians followed. Vascular plants made an appearance in the mid-Paleozoic, about 400 MYA. These plant pioneers had short, bare stems, but lacked roots and leaves. Cooksonia, the first known vascular plant, was about 5 cm high. Its leafless stems were topped with spore-filled caps. Some with woody stems, such as giant club mosses (e.g. Lepidodendron) and horsetails, became the first trees, up to 30 meters high. As Pangaea amassed, aridity became widespread in its interior, far from the weather ameliorating oceans. Coal Age swamps were supplanted by hardier vegetation such as the seed fern Glossopteris and early conifers replaced the drought intolerant swamp plants by the close of the Permian Period, 240 MYA. Larger plants included the Gymnosperms, such as cycads and pines, Glossopteris, and tree ferns, all good tree-top dinosaur browsing fodder. As Pangaea grew in diameter, its interior became increasingly arid.
Around 250 MYA, Pangaea began to break up into an upper or northern section, called Laurasia, which consisted of hunks of what was to be assembled as North America, Europe, and Asia. The lower section was called Gondwanaland and contained the land masses of Africa, South America, India, Madagascar, Antarctica, and Australia. The Laurasia-Gondwanaland split occurred before the time of flowering plants. Thus we find no early fossil Angiosperms in Laurasia but do find plants such as the extinct seed fern Glossopteris, the still extant Ephedra, and early reptiles common also to Gondwanaland.
According to the plant fossil record, Angiosperms had appeared by 125 MYA. Perhaps they started on a bank in the interior as a scrappy plant capable of rapid germination in a season and double fusion in the embryo sac after pollination. [In double fusion, a pollen grain contains two sperms. With the ova, one forms a zygote and the other combines with a nucleus to form the nutritious endosperm that sustains the seedling. Gymnosperms take two years to pollinate and their ovaries wastefully develop nutritious endosperms even if not pollinated. Often being wind pollinated, they tend to grow in proximity within groves.] Insects were attracted to Angiosperm flowers and inadvertently transported pollen, a more satisfactory and efficient method than the hit and miss of Gymnosperm windblown pollination and the water media pollination of other plant life. The Angiosperms were a success, rapidly evolving and spreading, radiating into many orders. By 90 to 100 MYA most modern orders had evolved. It is speculated that the Monocot sub-class (palms, grasses, aloes, orchids, bromeliads, lillies, etc.) evolved from an early Dicot (most other Angiosperms) and was possibly a water plant.
While the idea of continental drift is now established, its blessing from the scientific community came slowly. From 1620 onward, ideas such as the splitting of Africa and South America by the Biblical Flood were espoused by many, including the scientist Sir Francis Bacon in 1620 and Alexander von Humboldt, the pioneer explorer of the Americas in 1800. Another prevalent view until recently was that the appearance of the two continents fitting together was an illusion, and that the passage of plants and animals was facilitated by land bridges, now flooded. Antonio Snider in 1858 came close to a modern construct by fitting the continents together in an attempt to explain the occurrence of worldwide coal and fossil plant belts. More on this later.
Wegener And Plate Tectonics
Alfred Wegener, a German meteorologist and explorer, was the pioneer discoverer of the modern theory of continental drift (Figure 1). After wondering why tropical ferns had grown in London, Paris, and even Greenland, and why glaciers covered Brazil and the Congo at the same time, he authored several papers and four editions of his book The Origins of the Continents and Oceans, first published in 1915 (Reference 1). The evidence from his work drew not only from the study of ancient climates but also brought to bear evidence from a wide range of sciences. Wegener argued against the idea of land bridges between continents, but this view maintained its adherents. One sees what one expects to see! His detractors can today be passed off as ultra-conservative diehards but at that time it was not yet understood how continents could move. The sea-floor spreading mechanism for continental drift was not discovered until the 1970s. [ Darwin and Wallace had a similar problem. While their theory answered many questions, the mechanism of inheritance was unknown to them.]
In 1928 Wegener decided that a fifth revision of his book was beyond him. The relevant literature had become too extensive and specialized for a single worker to survey. In 1930 Wegener died during an expedition on the Greenland icecap. Before and after his death he and his work aroused passions. He was called a faker and mountebank, and in meetings called to discuss continental drift he was derided for playing fast and loose with data. George Gaylord Simpson, a noted vertebrate paleontologist, was amused to note that only one paleontologist at a meeting was sympathetic to the idea of drift, but that the drift proponents claimed that the paleontological data supported them. He commented in the American Journal of Science that "It must be almost unique in scientific history for a group of students admittedly without special competence in a specific field thus to reject the all but unanimous opinion of those who do have such competence" (Reference 2).
Wegener's opening line in his book was a statement of his philosophy: "Scientists still do not appear to understand sufficiently that all earth sciences must contribute evidence towards unveiling the state of our planet in earlier times, and that the truth of the matter can only be reached by combining all the evidence." To his detractors who recommended that his work be "thrown into a waste can," I would add that the essence of science is not only to find answers but to ask questions.
It is now realized that continents are not free to drift but are rigidly attached to plates which are moved by sea-floor spreading. A new term, Global Tectonics, covers the broad field of the deformation of the Earth's surface, including continental drift, sea-floor spreading, and transform faults and their relationship to earthquake shock waves.
The Breakup Of Pangaea
In the period from 100 to 80 MYA, Gondwanaland began to come apart. India separated, then Antarctica/Australia, and Madagascar. We know when and how this happened from a variety of sources including geology, botany, biology, radiochemistry, oceanography, and paleontology. Progress in putting this picture together over the past 20 years has been extraordinary. This activity had its beginning with Alfred Wegener, who gathered what he thought was overwelming evidence in support of his ideas on continental drift (Reference 1). He proposed that from 450 MYA to about 100 MYA most of the Earth's land was joined into a supercontinent which he called Pangeae. [Of Greek derivation. Pan Gaia means all Earth.] His 120 MYA Pangaea construct resembled the amalgam shown in Figure 2. Evidence for this included the vein of pre-Cambrian iron deposits which laced through what was to become Europe, Asia, North America, South America, Africa, India, Australia, and Antarctica; abutting geological provinces between the continents which matched when reassembled; and worldwide matching tin belts, as shown in Figure 3. Furthermore, the sequence and timing of The Breakup could be deduced from examining plant and animal fossil sites.
Continental drift is not a mere cruise control for continents. Without its collisions between plates, the raising of mountains and the altering of landscapes would be reduced, slowing evolution. Without the subduction of heavier oceanic plates under continental plates, the Earth would become a dead planet. Nutrients would pile up on the sea bottoms: carbon in the form of carbonates from atmospheric carbon dioxide; calcium carbonate from sea life; soil minerals and top soils flushed from land. Carbon dioxide is a major greenhouse gas. With its gradual loss, the Earth would grow frigid. As a result of subduction, the sea-floor sediment accumulation is thrust under land and eventually spewed out through volcanism.
Several locations have been proposed for Madagascar in the late Cretaceous Period (about 140 to 80 MYA). These include fits to Zanzibar at Tanzania, Beira at Mozambique, further south at Parfuri, or even offshore during the Cretaceous (Figure 4 and Reference 2). In my opinion, the evidence doesn't support Madagascar being offshore during the Cretaceous for several reasons: the good coastal fit at Beira; its isolation during the rise of the Angiosperms, which would interfere with its obviously being introduced to and inoculated by African plant families such as Didiera. From its unique flora and fauna, we would agree with the geologic estimates of its isolation from the African Mainland before the end of the Cretaceous. A tectonic map shows that the Madagascar geologic belts best match either Beira or nearby Parfuri, either of which seems reasonable. Since Beira at least superficially best fits the coastline, I have adopted it as the more likely site. The Zanzibar site was based on evidence available in 1930. Later work by Henri Besairie, summarized by John Wellington (Reference 3), found the Madagascar rough crystalline and the Karoo system sedimentary zones as a good geological and shape fit to the Mozambique plain.
The reasoning behind the idea of an offshore Madagascar, stationary in its present position, may have originated from the rapid travels of the India plate and its continuing pressure into Asia, forcing up the Himalayas. One might ask, "Shouldn't Madagascar, if it had a similar microplate, barge off vigorously?" Not necessarily. It could have been shoved off by an upthrust from a spreading center or, if on a plate, its movement could have been arrested by tectonic subduction.
Evidence For Pangaea
Evolution in biology and geology involves the flow of events. This includes changes with time and unexpected events along the way. A living plant or animal just happening to be in a certain place as these changes occur may have its future development determined. Taking the example of xerophytes, over 20 plant families in all the continents faced similar arid climatic conditions and evolved somewhat similar solutions. This is called convergent evolution. While this sounds obvious, as recently as the 1970s biogeography was controversial. Perhaps it shouldn't have been, given the work of Wegener, but there were still doubts about the idea of continental drift. This issue was settled by the discovery and measurement of sea-floor spreading in the 1970s, which confirmed continental drift as the driving mechanism. While there are problems to be worked out and details to be filled in, the continental drift theory has helped explain many biogeographic mysteries. I offer a few examples.
A vast coal belt traverses eastern North America, Europe, Asia Minor, and China. In Carboniferous times this belt followed a great equatorial circle: fossil peat bogs from low-lying swamps made of tree ferns such as Pecopteris, twining ferns, plants with stem-born blossoms (cauliflory). Fossils of the extinct seed fern Glossopteris are found preserved within coal seams in Africa, South America, the Falkland Islands, India, Afghanistan, Kasmir, northeast Iran, Northern Russia, Antarctica, and Australia. Its occupation of South America, Africa, India, and Australia is explained by their being contiguous in the Carboniferous Period, some 300 MYA, as depicted in Figure 5a, being dispersed in a 4-way split (Figure 5b). The Asian Glossopteris possibly spread from India.
Lystrosaurus was an early reptile with an early extinction. Its fossils have been found in southern Africa, Antarctica, and India, as sketched in Figure 6, also suggesting an early separation. Many other connections have been made by keen observers in both the field and laboratory. A classic was the Wallace line drawn by Alfred Wallace between Borneo/Southeast Asia and Celebes/New Guinea/Australia. Even though Borneo and the Celebes are neighbors, their biota and by surmise their origins are rooted in Asia and Australia, respectively.
There is an interesting example from a laboratory study of frogs and marsupials by Maxson et al. (Reference 4). Albumin proteins were found to act as an evolutionary clock. The tree frog subfamily Hylinae is distributed in the New World and Australia, but not Africa or Southeast Asia. This includes the New World genus Hyla (our many tree frogs) and the Australian Litoria (such as White's Tree Frog, now seen in pet stores). Since South America and Australia were connected by Antarctica, their ancestors must precede the separation of these two continents. The date arrived at for their geological separation was 70 MYA. This is at least 20 MY after Madagascar and South America separated from Africa. Picture, then, this separation sequence: India, Madagascar, and South America/Antarctica/Australia (with the Celebes and New Guinea group); then South America from Antarctica/Australia. [The frog fossil record begins with froglike Triadobatrachus (batrachus= Greek for frog) in the early Triassic, 220-230 MYA. It had a wide flat froggy skull and a tail. About 150 MYA, fossils were clearly recognized as frogs, forming their present order of Anura. They were placed in the sub-order Archeobatrachus and their surviving descendants were placed in Mesobatrachus, whose fossil record begins about 150 MYA. Mesobatrachus includes Pipidae, Rhinophyidae, Pelbatidae, and Pelodytidae. All other frogs, the new frogs, called Neobatrachus, were evident 50 MYA and include Hylidae and Litoria, mostly arboreal tree frogs, occurring in South America, North America, the West Indies, and Australia, but not Africa.]
Another interesting insight from Maxson involves marsupials, which occurred only in South America and Australia. The Antarctic landlink between these two continents is the same as that described in the frog albumin study and a marsupial albumin clock study also found that the geological separation occurred about 70 MYA. Even the marsupial parasites on the two continents are the same. [From fossil evidence it is believed that marsupials arose in North America in the Jurassic period. Placentals appeared in the late Jurassic or early Cretaceous. By the end of the Cretaceous, marsupials were in retreat, being supplanted by the placentals. There were several waves of placental advances: at the end of the Mesozoic, in the early Oligocene, and in the Miocene. Each wave displaced previous inhabitants. Marsupials reached Australia prior to its separation, leaving the placentals behind.]
From work such as the Maxson study, I would surmise that 70 MYA, at the time of the Australian-South American separation, Cactaceae had not yet evolved from South American Caryophyllales.
Cactaceae Comes To The New World
Angiosperms (flowering plants) originated in Africa about 125 MYA. The Angiosperm taxonomic order Caryophylalles is a monophyletic order which includes the closely related Cactaceae (cacti), Aizoaceae (Mesembs), Portulacaceae (Portulaca, Anacampseros, and Talinum), and Didieraceae (Alluadias and Didieras) from the New World, south and southwest Africa, and Madagascar, respectively. This suggests their origin from a common ancestor somewhere in southwest Africa prior to the Gondwana break-up. Both Cronquist and Rodman (Reference 5) placed the Mesembs as a sister family of Cactus (Figure 7a), but a recent DNA study by Hershkovitz and Zimmer (Reference 6) distances its kinship (Figure 7b). This suggests the scenario that the ancestral lineage of the Portulaca-Cactus-Didiera complex was split between Africa, the New World, New Zealand, and Madagascar. Hershkovitz and Zimmer estimated that cacti originated some 30 MYA (mid-Tertiary) with a later Tertiary diversification coincident with development of the American deserts.
The southern tip of South America is usually pictured as straight and southward pointing (Figure 8). With that arrangement, the ancestors of Cactaceae would have had to cross what was to become Africa in order to emigrate to South America. Wouldn't it be convenient if South America's continental shelf curled around the south end of Africa, as shown in Figure 5a, and abutted against what was to eventually break off and become Madagascar? Then the closely related members of Caryophylalles would have a point to radiate from and what was to become Cactaceae could inoculate nearby proto-South America without having to pass through a continent of varying climate zones. Is there evidence for such an arrangement? I believe there is. In Figure 8, note that the submerged South American continental shelf hooks to the east. It is my opinion that this shelf hooked around the Cape in an arc, abutting Mozambique and Madagascar, as shown in Figures 5a & 5b. Instead of having the ancestral precursor of cacti crossing the width of southern Africa to reach what was to break off and become South America, I believe it more reasonable and an easier trip to connect with the surmised continental shelf. I am suggesting that the ancestors of Cactaceae didn't go to South America but that South America came to it. To the best of my knowledge, there is no evidence against the Cape land arc. Confusion regarding this may have risen from uncertainties regarding the true continental edge of eastern Africa's coastline. This is not accurately known, land mass from the edge being found offshore on the continental shelf below 2,000 meters depth. [When continental plates split at a seam, the seams stretch and slump into a tapering shelf whose extent is usually estimated at being at a 2,000 meter depth. This works well for west Africa but is uncertain for the east coast.] I conclude from the existence of a southern and eastern shelf of South America, the uncertainty of the east Africa ancient coastline, and the possibility of a near Mozambique radiation center that East and South Africa, Madagascar, and South America were in proximity about 100 MYA, as shown in Figure 5. Interestingly, the Snider reconstruction of 1858 (Figure 9) aimed at explaining the occurrence of the worldwide coal/fossil plant deposits follows this pattern (References 2 & 7).
Another arrangement for the Pangaean split, similar except that Antarctica would nest between eastern Africa and South America, is a possibility. This arrangement, shown in Figure 10, would not alter the argument that those Caryophyllales which dispersed into the New World radiated from Mozambique, only that they would have had to traverse the territory now known as Antarctica to get there. This traverse could have occurred either before the Africa-Antarctica split or afterwards, when Antarctica was dangling at the tip of South America. The tip-dangling scenario appears to be the route taken by the Hyla and Litoria tree frogs and the marsupials, as previously discussed.
About 90 MYA, what was to become India had already separated from Gondwana and was on a collision course with the Asian mainland, due to arrive 20 MYA. About 80 MYA, South America was connected by Antarctica to Australasia; Africa and Madagascar were separated by water. Australasia separated 50 MYA. cacti's ancestors were in the proper place and at the proper time for a journey to the New World.
Collisions of the East Pacific Plate against the South American Plate 40 to 30 MYA raised the Andean Cordillera, producing volcanic activity which invaded giant batholiths, washing prodigious amounts of silt, sand, ash, and boulders eastward as far as the Amazon and Orinoco basins and the Argentine Pampas. A subsequent north-south compression further raised the Andes. These orogenies have caused and continue to cause earthquakes, rifting, and a subsiding of valley floors. The quickened movements of evolution almost always accompany great geological upheavals and climatic change. In this landscape cacti evolved. Those isolated on the east and west are distinctly different (Figure 11). The Eastern branch contains Austrocactus, Pyrrhocactus, Brasilicactus, Parodia, Wigginsia, Eriocactus, Notocactus, Blossfeldia, Gymnocalycium, Brachycalycium, Weingartia, Oroya, and Soehrensia. The Pacific branch includes Neoporteria, Copiapoa, Rodentiophila, Neochilenia, Horridocactus, and a few others.
Cactus Travels Through Time & Space
By Stan Yalof
At the Palomar C&SS 22 January 2000 meeting I gave a brief talk on a big subject. The talk marshaled evidence from geology, paleontology, field biology and molecular biology to identify the ancestral homeland of the cactus family in Panagea near Mozambique. Some of my conclusions built on the ingenious geology of Alfred Wegner, who conceived the theory of continental drift, as well as others referenced in the article printed last month in the Cactus Courier. I also discussed the timeline for cactus evolution and its transformation as they moved northward from South America to North America. About last month's article:
It covered only half of my talk, describing how the ancestor of cactus came to arrive on the South American mainland from Panagea.
The balance of the talk, from diversification due to the rise of the Andes Cordillera and the travel northwards to the USA was drawn from Part II of the story, still unpublished.
The illustrations for Part I were omitted last month due to a lack of space. They and their captions follow. Please refer to the text in last month's Cactus Courier for their elaboration.
The three-way split of Africa, Madagascar, and South America (with Antarctica and Australia still attached) occurred about 90 MYA. Members of the order Caryophyllales were separated within this split, evolving in their separate territories into mesembs, Didieras, and cacti. About 50 MYA, Antarctica and Australia separated from South America. From the Hershkovitz and Zimmer study, Cactaceae would not evolve from their Caryophyllales precursors until 30 MYA.
If, as Hershkovitz and Zimmer convincingly argued, cacti evolved 30 MYA, about 60 MY after South America became isolated from Pangaea, we are led to wonder what form cacti's ancestors took in this 60 million year stretch. Hershkovitz and Zimmer have provided a trail to follow. From their DNA work they have identified Talinum caffrum, a South African plant, as the base plant for a number of Caryophyllales species, including Cactaceae. Figure 12 summarizes this aspect of their work, which showed that early cacti were three branches distant from T. caffrum.
There is still the question of what triggered ancestral plants into becoming cacti some 30 MYA. Could the rise of the Andean Cordillera be responsible? According to the modern or systems view of Darwinism, evolution may not involve continuous gradual change. Species can remain constant for millions of years, as did human ancestors Australopithecus (Lucy) and Homo erectus. A recurring pattern in the history of Earth's biota is the period of growth and innovation following catastrophes. Such forks in the road, called planetary bifurcations, led first life and later life to great invention: photosynthesis; extraction of hydrogen from water; the use of potentially poisonous oxygen as oxidizer; the creation of lignin by early plants, making leaves, stems, and vascular systems possible; the seed ferns with their enclosed plant embryos (seeds) for protection and drought resistance; evergreen conifers, appearing 200 MYA during cold wet winters when glaciers appeared on several continents, wiping out the seed ferns; the Angiosperms, with their improved seeds and their cooperative arrangement with pollinators, appearing 130 MYA in response to increasing aridity; xerophyte species' physiological adaptations to extreme aridity. Cactaceae have developed an exceptional variety of adaptations to the xeric environment such as having a branchless, leafless, pleated, wax-coated stem with shallow roots and CAM/C-4 metabolisms. Primitive Cactaceae such as Pereskia aculeata are not suited to the xeric life, lacking the advanced adaptations. But they do have a feature shared with all other cacti, areoles. From the preceding, I would consider it reasonable to assume that the family Cactaceae was launched 30 MYA in South America, triggered by the rise of the Andean Cordillera. Its ancestors were Talinum or some further links.
Quite possibly, the ancestral species leading to cacti remained in stasis (not greatly changed) and in the same locality from 90 MYA to the Cordillera bifurcation. From classical Darwinism, one would expect continuous production of novelties and a gradual expansion of range. In practice, an expansion of range by new varieties into adjacent ranges is resisted either by a physical barrier or by entrenched species which occupy available ecological niches. To spread, a plant or animal species would have to displace some other biota from its entrenched niche. If this weren't the case, all species could continuously advance over time. Then the creatures in one place would not differ from others far off. A bifurcation is needed to overcome this ecological resistance. A catastrophe can be small: disturb the soil and weeds will take over. It can be medium-large: Krakatoa wipes an island clean of its biota and is recolonized from neighboring islands. It can be large: the raising of the Andean Cordillera, creating new climate patterns, geographic barriers, and opportunities for speciation for plants and animals that happened to be around.
Subsequent collisions of the East Pacific oceanic plate against the South American plate produced an era of Andean mountain building and landscape changes which separated the evolving Cactaceae into Eastern and Pacific sub-groups. In a subsequent brief contact with North America territory, I imagine that Cactaceae moved into southern Mexican territory and began to radiate northward. This move was abetted with the joining of North and South America several MYA, cutting off ocean circulation about these two continents, creating cold ocean uplifts on their Pacific coasts from the returning Arctic and Antarctic currents, thus producing increasing aridity. This aridity was intensified at 30 degrees north and south latitudes due to atmospheric circulation (Hadley Cells) and the presence of coastal mountain ranges. About 10,000 years ago, glaciers retreated and a warming trend began that boosted human agriculture and also led to further aridity in northern Mexico and western United States territories. Cactaceae had evolved into a superb xerophyte with CAM/C4 metabolism, water storage by stem and/or caudex, thick and waxy cuticles, reduction to a pleated stem, shielded stomata, high-temperature resistance, and quick-absorbing root systems. They expanded into this increasingly arid territory, evolving into new species. Biogeographic isolation within the Mexican mountain ranges, in the Baja California peninsula and its offshore islands led to new species and varieties, many recently evolved, many found nowhere else (endemic).
New research methods have extracted information from living systems that could not have been conceived of 10 to 20 years ago. Much of the past field and herbarium work has to be reexamined. To confirm that the catastrophic bifurcation of the Andean Cordillera triggered the evolution of cacti, the cacti mentioned in Figure 10 should be examined in all ways to determine their relationships. The New World Talinums should be further explored as to their links to cacti.
1. "The Origin of Continents and Oceans," Alfred Wegener, Dover Edition (1966).
2. "Continental Drift: The Evolution of a Concept," Ursula Marvin, Smithsonian (1973).
3. "South Africa, A Geographical Study," Vol. 1 Physical Geography, Chapter 15, PP 460-465; John H. Wellington (1955).
4. "Continental drift and the use of albumin as an evolutionary clock," Linda Maxson et al., Nature, Volume 255, No. 5507, PP 397-400 (29 May 1975).
5. "The Cactus Primer," A. C. Gibson and P. W. Nobel, Harvard Press (1986), PP 244-247.
6. "On the evolutionary origins of the cacti," Mark Hershkovitz and Elizabeth Zimmer, Taxon46 (May 1997), PP 217-232.
7. "La Création et Ses Mystères Dévoiles,"Antonio Snider-Pelligrini (1858).
Figure 11: Cacti split into two branches by the Andean Cordillera ______ Group 1 Lobiviae Sub-tribe 2 _______ Austrocactinae ________ Sub-group 1 Austrocactus Eastern branch Pyrrhocactus Brasilicactus Parodia Wigginsia Eriocactus Notocactus ________ Group 2 _______ Frailea Austroechinocacti Blossfeldia Soehrensia Oroya Gymnocalycium Brachycalycium Weingartia Sub-group 2 Rodentiophila ________ Pacific branch Neochilenia Horridocatus Delaetia Reicheocactus Neoporteria Eriosyce Islaya Pilocopiapoa Copiapoa Figure 12: The Cactaceae ancestral hierarchy evolving from Talinum caffrum (adapted from Hershkowitz and Zimmer). Didieraceae (Madagascar, 11 species) Basellaceae (Madagascar & pantropical, 11 species) Talinum caffrum ____ (S. Africa) Portulacaria (S. Africa, 40 species) Talinum paraguayense (S. Am.) Cactaceae --- Pereskia aculeata (N. and S. Am.) __________________ Maihuenia poeppigii (S. Am.) Perekiopsis porteri (N. Am.) Pereskia grandiflora (S. Am.) Figure 13: Cacti split into two branches by the Andean Cordillera, as for Figure 11, but with genera lumped according to a RBG (Kew) listing. Locations for the genera outside the original listing are: (1) N. Bolivia, S. Argentina, Paraguay, Uruguay, S. Brazil; (2) N. Arg., Bol., S. Peru, N. Chile; (3) N. Arg., NE. Bol; (4) W. Arg. from Rio Colorado to Jujuy and Salta. ______ Group 1 Lobiviae Sub-tribe 2 _______ Austrocactinae Austrocactus Parodia Brasilicactus Wigginsia Eriocactus Notocactus _______ Sub-group 1 Frailea Eastern Branch Blossfeldia Gymnocalycium Brachycalycium Weingartia ...................... Rebutia (3) Pyrrhocactus ................... Neoporteria (4) Soehrensia ...................... Echinopsis (1) Oroya .............................. Oreocereus (2) Neoporteria Neochilenia Horridocatus Delaetia _________Sub-group 2 Islaya Pacific Branch Eriosyce Rodentiophila Copiapoa Pilocopiapoa Reicheocactus .................. Rebutia Alfred Wegner