the cephalium in cacti

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Marlon Machado
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the cephalium in cacti

Post by Marlon Machado »

One thing that puzzles me is the evolutionary route the various cephalium-bearing genera have taken - both the columnar ones and Melos/Discos. What is the purpose of them, and/or how did they come about, do we know?

Stuart Estell
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Hi Stuart,

How and why the cephalium evolved are indeed quite interesting questions. This structure evolved several times in the family, in many independent groups both in North America and in South America. Thus, there must be a very good reason for so many different plants have evolved similar structures.

With a few exceptions, each areole of all cacti produce a single flower, and that is it - after this flower is produced, the areole become inactive. Exceptions to this rule are Myrtillocactus in North America, which can produce several flowers per areole:
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Myrtillocactus geometrizans. Photo: Gerard Ardisson's Cactus & Succulents Photo Gallery
and Neoraimondia in South America, whose areoles not only produce several flowers per areole but they also flower year after year, the areole growing each year and becoming longer, like a miniature cephalium:
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Neoraimondia gigantea. Photo: Mauseth's Lab
But these exceptions aside, what usually happens in cacti is that once a areole has produced a flower and fruit, it becomes inactive. So, in order to produce more flowers, the cactus has to grow more areoles. And to grow more areoles the cactus has to grow new stem. However, the growth of new stems can be a very costly effort - for each new few centimeters of stem, the plant has to produce lots of inner tissue, epidermis, etc. which demands lots of energy and nutrients. From a cost/benefit perspective, it can be very costly to produce flowers and seeds for reproduction if each time the cactus has to grow new stems and thus spend a lot of energy in the process.

Many cacti devised clever ways of minimizing the amount of energy spent for reproduction by producing more areoles in a smaller area, thus producing more flowers without having to grow significant amounts of new stem. Take for example Astrophytum myriostigma. Young plants of this species have areoles well-spaced along the ribs:
but as the plants grow older the areoles are produced at closer intervals, and big mature specimens have a continuous line of areoles along the ribs:
Thus, a mature Astrophytum myriostigma reaches a good balance where it can produce lots of new flowers each year without having to spend too much energy in growing new stem.

Another example is the Senita Cactus, Lophocereus schottii, which is a columnar cactus species from northweastern Mexico and southern Arizona, US:
Juvenile stems have well-spaced areoles:
Which in maturity will develop very closely spaced areoles, with long spines:
Cacti that develops a cephalium just took a step further on this concept of saving energy while maximizing reproduction potential. In cacti with cephalia, the reproductive parts of the stem are composed of very closely spaced areoles, so that many flowers can be produced with a minimal amount of energy spent with the growth of new stem. There are various types of cephalium:

apical cephalia like those produced by Melocactus:
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Melocactus zehntneri. Photo: Marlon Machado
Discocactus:
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Discocactus placentiformis. Photo: Marlon Machado
and Backebergia:
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Backebergia militaris. Photo: Mauseth's Lab
lateral cephalia like those produced by, among others, Cephalocereus:
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Cephalocereus senilis. Photo: http://www.astrobase.de/
Espostoa:
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Espostoa guentheri. Photo: Philippe Corman, http://www.cactuspro.com/
and Coleocephalocereus:
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Coleocephalocereus aureus. Photo: Marlon Machado
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Coleocephalocereus purpureus. Photo: Marlon Machado
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Coleocephalocereus buxbaumianus. Photo: Marlon Machado
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Gerard Delanoy with Coleocephalocereus goebelianus. Photo: Marlon Machado
and apical cephalia with recurrence of vegetative growth (ring cephalia) like those of Arrojadoa:
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Arrojadoa dinae. Photo: Marlon Machado
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Arrojadoa marylanae. Photo: Marlon Machado
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Arrojadoa rhodantha. Photo: Marlon Machado
and Stephanocereus:
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Gerard Delanoy with Stephanocereus leucostele. Photo: Marlon Machado
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Stephanocereus leucostele in fruit. Photo: Marlon Machado
Besides the energy-saving benefit, the cephalium also has another great advantage, which is to afford protection to the flowers and fruits while they are still developing. The areoles of the cephalium usually produce copious amounts of bristles and wool which protect the young flower buds until they are ready to open, and also the young fruits until they are ripe. The flower buds and young fruits are hidden within the cephalium, enveloped by the bristles and wool, and not visible until they are mature enough to appear in the surface of the cephalium.

This is a very interesting subject indeed, and a good place to find more information about the cephalium is the website of Prof. Dr. James D. Mauseth. His text is very instructive, but beware that he uses a lot of technical terms. And, if you still wants to learn more, there is an excellent review also published by Mauseth:

Mauseth, J.D. 2006. Structure?Function Relationships in Highly Modified Shoots of Cactaceae. Annals of Botany 98: 901?926.

Cheers,
Marlon Machado.

Institute for Systematic Botany, University of Zurich, Zollikerstrasse 107, CH-8008 Zurich, Switzerland.
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Stuart Estell
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Re: the cephalium in cacti

Post by Stuart Estell »

Marlon, thanks for such a detailed and lengthy explanation. Fascinating stuff.

Do you have any pictures of dissected cephalia showing the compressed stem growth beneath the bristles by any chance?
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Re: the cephalium in cacti

Post by North Lincs Mike »

Marlon

Many thanks for this the most detailed and authorative answer to any question I have seen in the few years I have been on the Forum.

Moderators - I am not really clear how the reference sections of the Forum work but it would be good if this and Marlon's Melo thread could be retaine for future reference.

Mike
Newish collector/grower; mixed lot in a new 12x18 g/h and my old re-erected 10x8 - good job I didn't sell it !
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Re: the cephalium in cacti

Post by Cactus Jack »

:notwo: Great series Marlon , very informative(tu)
Stephen.. Bangor. N. Ireland.
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DaveW
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Re: the cephalium in cacti

Post by DaveW »

One of the characteristics of Backeberg's Chilenia was that they often produced more than one flower per areole, most are now known as Eriosyce.

Beautiful pictures Marlon!

DaveW
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Re: the cephalium in cacti

Post by Susanne »

Hello Marlon

Thank you very much! This article was great. I read it with great interest. I also liked the photos. It is good to get so much information.:morng:
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Re: the cephalium in cacti

Post by Phil_SK »

I think Weingartia can do the several flowers per areole thing too.
Phil Crewe, BCSS 38143. Mostly S. American cacti, esp. Lobivia, Sulcorebutia and little Opuntia
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Re: the cephalium in cacti

Post by DaveW »

Some Weingartia's were the other species in Backeberg's Chilenia Phil.

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Re: the cephalium in cacti

Post by Nick »

Marlon - i second the above - great description. And thanks for the reference - it`s good to see a free one - they mostly come in around the $30 which is a bit much for Joe Public!
Nick
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Re: the cephalium in cacti

Post by Vic »

Yes a fascinating explanation Marlon just like attending a lecture, some great photos and amazing plants. I hadn't really ever thought why a cactus would produce a cephalium but it's makes great sense. I wonder why more South American cacti do compared to North American.
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