viviparous from the flower
The Origins of Viviparism
by Jorge Monteverde
Buenos Aires, Argentina
© 2009, all rights reserved
Translated from Spanish by Fernando Molina
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Photo credits with large images
One of the most common vegetative propagation mechanisms for
cultivated waterlilies in our water gardens is generally called
viviparism-- which generally refers to a living being
that produces live offspring, and in botany to those plants which
produce bulbs or seeds that germinate while still attached to
the parent plant-- and surely it is close to our experience since
a good number of these plants display such behavior as a reproductive
strategy. But we must keep in mind that these waterlilies, mostly
hybrids or cultivars, inherited this feature from one or more
of their ancestors. If we go back through successive generations,
somewhere along the line we are bound to find the link with the
species that originated this particular feature.
Among the approximately 52 known species, viviparous
behavior is pretty rare and those that display it habitually
can be counted with the fingers of one hand. In fact, only three
known species today have that reproductive form as a primary,
almost primary, or combined resource: Nymphaea micrantha
Gill. & Perr., Nymphaea lasiophylla Mart. & Zucc.,
and Nymphaea prolifera Wiersema.
What can be highlighted at first sight from these species?
All three are tropical.
Only one of them (N. micrantha) is a day-bloomer,
while the other two have nocturnal anthesis.
Among recognized subgenera, N. micrantha belongs
to Brachyceras while N. lasiophylla and N. prolifera
belong to Hydrocallis.
Their type of viviparism:
- N. micrantha with its gemmipary(1)
property (meaning the development of young plants from vegetative
buds borne on the parent body) generates new individuals only
from its leaves, developing epiphyllous plantlets (from
the Greek meaning upon a leaf).
- N. lasiophylla and N. prolifera do it from
their flowers, by means of abortive tuberiferous flowers
in a process called pseudovivipary(1)
by Sculthorpe in 1967 and vivipary (2)
by Wiersema in 1987.
Back from botany to our daily experience, as noted above there
are several viviparous cultivars. Among those we consider tropical,
their viviparism from the leaves corresponds, in ALL cases, to
their N. micrantha descent, namely N. Daubenyana,
N. Panama Pacific, N. Tina,
N. Lindsey Woods and many more.(3)
These waterlilies evidence their viviparism in a very pronounced
manner, since adult plants will make this mechanism their most
common form of propagation.
Stages of viviparism from the leaves
of tropical day blooming waterlilies
It is altogether different when we refer to hardies, where
in some this phenomenon is rare or almost unseen -- might even
never be seen -- but it exists. In others it is somewhat more
frequent. To summarize, we could say that those that present
this feature more or less markedly are slightly or eventually
viviparous and their reproduction does not depend on this particular
characteristic. Here N. micrantha is not involved, since
in this group of waterlilies in ALL cases there is flower
-- but NOT leaf -- viviparism.
We could then feel tempted to consider that viviparous hardies
have some kinship with N. lasiophylla or N. prolifera,
since only these species present vegetative propagation in their
floral structures as indicated above.
Is this possible? Lets say that almost anything is possible,
but it is not likely, because even though N. lasiophylla
was described in 1832 there are no references to its being used
in any hybridization. Furthermore, we know that they belong to
another subgenus, and intersubgeneric crosses have been known
and proved in more recent times (I admit, by the way, that this
is a rather weak argument). Let us add that the nonexistent horticultural
attraction of this species would make it unlikely for a professional
hybridizer to utilize it aiming to improve future generations
of cultivars. Lastly, we could consider as proof of its use that,
besides its being viviparous, its night-blooming characteristic
could have been transferred too -- but we do not have night-bloomers
among hardy waterlilies
Similar considerations might be added by specialists pro and
con, but ultimately no one can really attest or deny the N.
lasiophylla descent, which is why we subscribe the UNLIKELY
hypothesis given its more limited scope.
So if we discard N. lasiophylla and leave aside N.
prolifera for identical reasons, besides its being unknown
in a horticultural sense, we are left with nothing. So how is
it possible that we have flower-viviparous hardies?
Renowned experts say Colonel A.J. Welch was the first
hardy waterlily to sprout new plantlets from some of its flowers,
in small percentages, but sure enough. It is decidedly hardy,
viviparous, and the first to be known. So where did it come from?
There are some who think, among other considerations, that
N. Colonel A.J. Welch could be a non-described
species that became extinct in its native or natural state. Most
are led to believe, however, that Latour-Marliac (from whose
business it originated, although it cannot be said that he himself
hybridized it) or whoever actually made it used N. mexicana
and therein might lie the answer. This makes sense, because Latour-Marliac
used N. mexicana extensively and N. Colonel
A.J. Welch, given its color and its flowers above the water
level, surely carries its descent. Paradoxically, experts express
some doubts by noting that N. mexicana is not described
or known as viviparous from its flowers; it generates runners
or clusters in the shape of bananas in the root area.
In agreement with the above view, in my opinion N. mexicana
has caused this particular feature in viviparous hardies, including
N. Colonel A.J. Welch. To substantiate this
position, I offer the following evidence: 1) a scientific article
published in India in 1974 which explains this (4);
2) the testimony of Grant Mitchell, known for his love of lotus
and his trips to the Far East, reported extensively in our lists,
who some years ago in an IWGS forum said he had observed a flower
in N. mexicana with a little plantlet growing on it, and
lastly 3) the experience of a founding member of WGI, the now
famous hybridizer Pairat Songpanich, for having achieved the
almost impossible blue hardy called N. Siam
Blue Hardy. He has made use of N. mexicana for hybridization
and says it is a flower-viviparous waterlily based on his personal
experience. This feature, so far unique within the genus Nymphaea,
is certainly quite elusive, it does not appear in all N. mexicana
or is very weak, or else depends on very specific conditions,
and that is why it has largely not been observed.
This could have been the reason why using such a precise type
of N. mexicana with that particular feature to obtain
N. Colonel A. J. Welch gave its descent such
a strong trait, while it did not happen with many other hybridizations
past and present using the same species (N. Marliacea
Chromatella, N. Moorei, N. Aurora,
N. Andreana, N. Yellow Sensation,
Something similar could have happened with N. Colorado,
not reported as viviparous in books and other publications or
in general knowledge, but confirmed through personal comment
by the renowned hybridizer Rich Sacher in Louisiana USA, by Pairat
Songpanich and by the award-winning Dr. N. Nopchai Chansilpa,
who hybridized N. Mangkala Ubol, both from
Thailand. Also our good friend James Knock in Britain had already
reported this behavior in N. Colorado when
he wrote a memorable tribute to the work of Kirk Strawn in 2005
Strawn, who developed this cultivar in 1994, also developed many
others with similar characteristics, such as N. Barbara
Dobbins, N. Georgia Peach and N.
Clyde Ikins among others, using N. mexicana
as a pollen parent.
N. Barbara Dobbins, N. Cherokee,
N. Georgia Peach, N. Perrys
Red Star, N. Perrys Pink Delight,
N. Perrys Viviparous Pink and N.
Phoebus are the better known cultivars showing this
feature. Based on this new review, we can add N. Clyde
Ikins, N. Colorado, N. Innerlight,
N. Lemon Mist, N. Mangkala Ubol,
N. Patio Joe, N. Peace Lily,
N. Peachglow, N. Steven Strawn,
N. Texas Dawn and, surprisingly, N.
Lily Pons and N. Helvola.
It should be noted that the names of cultivars mentioned here
displaying this feature, even if observed only once, were provided
by some of those for whose help I thank at the end of this article
and who responded to my queries.
There are, however, some occurrences that seem to be exceptions
for the time being. Again, Pairat Songpanich tells us about some
of his hybrids, all of them odorata rhizome types
without apparent kinship with N. mexicana, that have also
shown this behavior. Such are the cases of N. Mayla
x N. Perrys Fire Opal, N. Perrys
Fire Opal x N. Sirius, and (N.
Perrys Fire Opal x N. Sirius)
x N. Sirius.
My initial working hypothesis before writing this article
was trying to prove the incidence of N. mexicana in this
type of viviparism. I believe this objective has been achieved,
although there are still some obscure areas which leave room
for further learning or understanding about this exceptional
behavior. Later its scope widened to cover all hardies, and finally
it came to include all the lesser known aspects of the family.
With respect to the subgenus Lotos, there are two scientific
reports, one book and several personal testimonies concerning
this asexual reproductive ability:
1) Barber, C.A. (1889), On a change of flowers to tubers
in Nymphaea lotus var. monstrosa, Annals
of Botany IV (13): 105-116.
2) Mitra R.L. and K. Subramanyam (1982), Is Nymphaea
rubra Roxb. ex Andrews an apomict?, Bulletin of
the Botanical Survey of India 24: 83-86.
Perry Slocum also assigned flower viviparism to N. lotus
in the description of this species in both editions of his book,
An Encyclopedia of Water Lilies and Lotuses (6).
We should also add the personal comment by the renowned hybridizer
Craig Presnell from Florida, who confirms the presence of flower
viviparism in N. Missouri of N. lotus
descent; the report by Dr. N. Nopchai Chansilpa who attests to
having seen several offspring of N. lotus with this behavior;
Nan Baileys report from Australia on an unnamed night-blooming
hybrid; and a report from my own country, Argentina, on N.
< ^ N. 'Missouri' >
Almost to finalize, a significant reference to subgenus Anecphya
will allow us to be truly original by reporting for the first
time ever a case of flower viviparism in N. gigantea.
Australian WGI member Roger Bagley witnessed this phenomenon
some years ago in the northern coast of New South Wales, more
precisely in Glenugie Creek, 15 km (9 miles) south of Grafton.
He scanned an old photograph taken in the field to illustrate
the phenomenon, even though it is not too visible because of
This report on Anecphya is the first and only one to
date. Andre Leu, an unavoidable reference and keen specialist
on this subgenus, refers to this case as an oddity of nature.
Even considering it exceptional, as it surely is, the ability
to propagate in this manner indeed exists.
Before we came upon this information, such asexual propagation
by abortive tuberiferous flowers appeared to be related only
to subgenera around the syncarpiae group. Dr. Wiersema
thus indicated in his description of N. lasiophylla:
rare occurrences of similar structures have been reported for
other syncarpous Nymphaea, and suggested reading
Moham Ram et al (4) and Mitra and Subramanyan
on N. rubra.
A separate paragraph must be dedicated to N. Jongkolnee,
the ancestral, mysterious, emblematic waterlily from Thailand.
Tropical and day-blooming, this exotic waterlily has lost its
ability for sexual reproduction, partly surviving also through
this mechanism of floral viviparism. However, until its taxonomic
status is revealed (species, natural or artificial hybrid, mutant,
etc.), we shall not include it in any subgenus and it will remain
apart waiting for further future research. If eventually, as
many believe should be the case, it came to be included in the
subgenus Brachyceras, it would be an exception and would
complete the circle within Apocarpiae, each subgenera
having one representative with this feature.
As a statistical synthesis, we can state that we ended up
with flower viviparism events in both groups (syncarpiae
and apocarpiae) and in four (Nymphaea, Hydrocallis,
Lotos and Anecphya) of the five or six admitted subgenera,
and with leaf viviparism in one group and one subgenus (Brachyceras).
As a result, each of the familys taxonomic division or
subdivision has at least one representative of viviparous behavior.
We should also point out that of the 19 waterlily names included
in this report (not including N. Colonel A. J. Welch),
10 are cultivars obtained by Kirk Strawn. Considering that some
of his other cultivars besides the above-mentioned have a similar
genetic descent, with N. mexicana as polen parent, it
would not be surprising if one of these might be added to the
list in the future when such an eventual occurrence is observed
and testified by a fellow water gardener.
One final comment: the most amazing reports came from the
Orient, mainly on N. Helvola and N.
Lily Pons and on odorata without the
alleged intervention of N. mexicana. It seems this phenomenon
is less observable in the West, while in Thailand with Pairat
Songpanich and Dr. N. Nopchai Chansilpa, and in China with the
great hybridizer and lotus and waterlily specialist Prof. Huang
Guozen of the Quingdao Biological Institute (Quingdao Water
Lily World), they see it as a rather common behavior.
I am grateful to Roger Bagley, Nan Bailey, Mauro Bergamo,
Paula Biles, Dr. N. Nopchai Chansilpa, Alani Davis, Daniel Flouret,
Huang Guozen, Don Hamburger, Harro Hieronimus, James Knock, Andre
Leu, Carlos Magdalena, Asta Miklius, Walter Pagels, Craig Presnell,
Andreas Protopapas, Rich Sacher, Pairat Songpanich, Martin Sturge,
Charles B. Thomas and Werner Wallner for their participation.
I want to express my special gratitude to my friend Dr. Jaime
Raúl Bonilla Barbosa, a professional specialist in aquatic
botany at the Autonomous University of the State of Morelos,
Mexico, who made a technical and formal revision of this article.
1) Sculthorpe, C.D. 1967. The Biology of
Aquatic Vascular Plants (London: Edward Arnold) 610 p.
2) Wiersema, John H. 1987. A Monograph of
Nymphaea subgenus Hydrocallis (Nymphaeaceae). Systematic
Botany Monographs 16: 1-112.
Waterlilies - www.victoria-adventure.org
4) Mohan Ram, H.Y., Laxmi Nayyar, Vijay (Department
of Botany, University of Delhi).1974. A Case of Reversion
of Flower of Nymphaea mexicana to Vegetative Condition,
Current Science 43 (9): 291.
Knock's Tribute to the Brilliance of Kirk Strawn - www.victoria-adventure.org
6) Perry D. Slocum, Peter Robinson and Frances
Perry. 1996. Water Gardening, Water Lilies and Lotuses