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Porophyllum Saxifrages riddles

Hans Meulenbelt
Adapted from a talk given at the Saxifrage 2016 Conference at Waterperry

I would like to start by quoting Reginald the words ‘genos’ (genus) and ‘eidos’
Farrer: ’In no great family is there more (species) in his works that were the first
fearful confusion as to names; some to rank the world’s organisms. After the
Oedipus is sadly needed to take the race Greeks, Carl Linnaeus was probably the
in hand and unravel its riddles’. next person to organise out thoughts
about the taxonomy of plants. Linnaeus
When we deal with Porophyllum thought of species as constant things
Saxifrages we certainly have a lot of that did not change in time. However, as
riddles to solve, wrong names on labels we all know, the well-known European
for sure but also in understanding species Saxifraga burseriana shows a
the species. This quotation appears at lot of variation, not only in the rosettes
the start of Dr Radvan Horny’s book but also in the flowers. But still we all
on Porophyllum Saxifrages. A book he recognise these plants as belonging to
devoted twenty years of his life to. He that species.
died last Christmas and so this lecture is Charles Darwin suggested that species
in honour of his legacy. Our thanks also were not constant but changed in time.
go to John Byam Grounds, because he This idea also influences the way we
made it possible that the English version look at taxonomy. The way that plants
of this book was published. are classified today represents just a
moment in time. It does not take account
of everything that has come before or
will happen in the future.

In search of an answer to the question


‘What is a species ?’ I stumbled upon the
Nature Education website and found
three definitions. There is a biological
species concept that defines a biological
species as a group of organisms that can
reproduce with one another in nature
and produce fertile offspring. Species
1 are characterized by the fact that they
are reproductively isolated from other
When thinking about the subject of groups, which means that the organisms
this lecture I wondered what exactly in one species are incapable of
constitutes a species. The concept goes reproducing with organisms in another
all the way back to Aristotle who used species. This concept works well for

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animals but not for plants because they Take a look for example at Saxifraga
often easily hybridise. aretioides. Not so long ago we
understood the plants growing in the
Taxonomy is a scientific system that Picos as belonging to this species. But
classifies organisms into categories now we consider them to belong to a
based on their biological characteristics. different species, Saxifraga felineri. So
In this biological classification system even in Europe, where plant life has been
the term species defines the lowest studied for centuries, our understanding
taxonomic rank. This second definition of Saxifrage species is changing.
is a little like what we do when we put a
label in a pot! We decide on a name. Today we grow many Porophyllum
species in our gardens and, of course,
Species can also be defined based on a huge number of hybrids. The story
a shared evolutionary history and of the Porophyllum hybrids starts with
ancestry. This method of defining species a publication written by D. Dewar in
is called phylogenetics, which is the 1890. In this article Mr Dewar draws
study of the evolutionary relationships attention to Saxifraga Boydi, now known
among organisms. The evolutionary as Saxifraga x boydii ‘Old Britain’. He
process by which a new species comes writes that this hybrid “was supposed to
into being is called speciation. be a cross between S. Burseriana and S.
Rocheliana”. I don’t know why as it has
So there are different ways of defining yellow flowers. It was Mr Dewar who
what makes a ‘species’. pointed out to Mr Boyd that S. aretiodes

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was more likely to be involved. Probably but when I show pictures of these plants
the first riddle solved. I am always told that they are probably
not the real thing because they look
Saxifraga ‘Old Britain’ is the parent far too healthy. Everyone states that S.
of the cultivars S. ‘Cherrytrees’ and lilacina cannot be grown in compost
S. ‘Faldonside’. In S. ‘Cherrytrees’ containing lime and must be grown
the rosettes are much closer to S. in an acid compost. However, in the
aretioides, while the flowers are closer original article by Walter Irving there is
to S. burseriana, but in S. ‘Faldonside’ no mention of this and he states that the
the rosettes are close to S. burseriana plant was grown in a ‘normal compost’
and the flowers closer to S. aretiodes. So at Kew. Reginald Farrer again makes
it’s always something of a surprise what no mention of this and only stipulates
you get from a combination of species. that it should be given shade. The plants
that I have as S. lilicina are grown in my
A huge development in the history of normal compost and they grow well in
the Porophyllum Saxifrages happened this mixture.
when a guy from India sent seed to Kew
Gardens from which a few seedlings After the introduction of this new
were grown and these plants were used species from the Himalaya it did not
to describe a new species by Duthie in take long for the first hybrids to appear.
1904. Of course this is Saxifraga lilacina. The first one was Saxifraga x irvingii
It was first mentioned in literature ‘Walter Irving’ (1909), a cross between
in an article, also in 1904, by Walter S. burseriana and S. lilacina, although
Irving. According to Irving the seed was this was not recognized by Walter Irving
collected in the Kumaun but according at the time. It was Dr Radvan Horny who
to Mr Duthie the seed was collected in solved this riddle.
Hazara in what is now Pakistan. The
whole history of Porophyllum Saxifrages An important next step in the history
is just full of riddles! of the development of Porophyllum
cultivars was the introduction of
If you look at the famous publication by several wild collected hybrids. Franz
Harry Smith (Saxifraga of the Himalaya I Sündermann received plants from Saint
Section Kabschia - 1958) you won’t find Beat in the Pyrenees. He had ordered S.
a description of Saxifraga lilacina - it aretioides and S. media, but among the
is only mentioned in the key. I did find plants he received he recognised some of
a description of S. lilacina in the digital them to be hybrids of these species. One
flora of Pakistan a few years ago, but for of these he describes as S. lapeyrousii.
some reason it is not mentioned at all in This particular form was used by
the current online version. It has always Reginald Farrer to create S. x anglica
been a mystery plant, which is strange if ‘Myra’ (1918). What surprises me, and
you realise that this species is involved this goes for Myra and all the other S. x
in so many of our present day cultivars. anglica cultivars, is that although one of
I grow plants that I think are S. lilacina, the parents has yellow or orange yellow

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flowers, there is no trace of yellow in the godroniana. This was used extensively
offspring. by Russell Vincent Prichard to raise
many different S. x anglica cultivars.
Now that S. lilacina was available as Many of the early ones are now lost or
a parent many new possibilities had wrongly named.
arisen. Crossing it with Saxifraga
stribrnyi gave us S. x hornibrookii Another riddle solved by Radvan
‘Riverslea’ (1925). There are many more Horny concerns the parentage of the
and all are strikingly beautiful. Another cultivar ‘Miss Gertie Prichard’ (1928).
early cross made by Sündermann was It was considered to be of S. x anglica
S. x kellereri ‘Suendermannii’ (1915). parentage, but Horny proposed S.
I don’t understand why this plant is burseriana to be one of the parents, so
not offered more often as it does really it became a S. x megaseaeflora cultivar.
well outside in the rock garden and is
extremely beautiful. Sündermann was Probably the most important plant in
also responsible for describing another the history of Porophyllum cultivars
notomorph, S. luteo-purpurea nm is S. x anglica ‘Winifred’ (c. 1935). It
was raised from an S. x anglica cultivar
(maybe S. ‘Cranbourne’) and S. lilacina
and the result was one of the most
beautiful Porophyllum cultivars. As a
giant bonus it produces seed readily and
has fertile pollen, making it possible to
use it as a parent. The list of cultivars
raised using ‘Winifred’ is almost endless

1 - Karel Pech - J.Byam - Radvan Horny


2 - Saxifraga x boydii
3 - Saxifraga anglica ‘Winifred’
4 - Saxifraga x megaseaeflora ‘Karel Čapek’
5 - Saxifraga x kellereri ‘Suendermanni’
3

4 5

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and these cultivars show how the colour almost stemless flowers. Karel Lang has
spectrum has increased using it. Just used it to produce S. [Exclusive Group]
look at the names of this ‘Winifred’ ‘Excellent’ (2006), using S. lilacina as the
offspring and you will realise how other parent. This cultivar is much more
important this cultivar has been. S. x colourful than either parent. Another
megaseaeflora ‘Karel Čapek’ (1975)’, example of a cultivar with big almost
S. x goringana ‘Nancye’ (1987), S. x stemless flowers is S. [Literary Group]
poluanglica ‘Red Poll’ (c. 1988), S. x ‘Johann Wolfgang Goethe’ (2006). We
siluris ‘Joachim Barrande’ (1998), S. did not have that before we used S.
x anglica ‘Amberglow’ (1999) and S. lowndesii.
‘Morava’ (1993) all were raised from S.
‘Winifred’ and all of them belong to my The introduction of Saxifraga dinnikii
favourite cultivars. No wonder it is still and Saxifraga columnaris, both species
being used by hybridizers today. from the Caucasus, has also added new
possibilities for hybridising. S. dinnikii
One other species that has opened up has huge flowers and the crystalline
new doors is Saxifraga lowndesii. One structure of the flower has given an added
of the earliest attempts to use it was by quality to our cultivars, for instance in S.
Oldrich Maixner and Jan Bürgel. They [Noble Group] ‘Moulin Rouge’ (2006),
created the cultivars that belong to the raised by Jan Bürgel using S. lilacina as
Blues and Beat Group. One added feature the other parent . Karel Lang crossed it
is that it has brought us cultivars with with S. aretioides to create S. [Superlativ

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10

7
6 - Saxifraga x poluanglica ‘Red Poll’
7 - Saxifraga x siluris ‘Joachim Barrande’
8 - Saxifraga ‘Excellent’
9 - Saxifraga x anglica ‘Amberine’
10 - Saxifraga x dinninaris ‘Ray Woodcliffe’

Group] ‘František Holenka’ (2010),


adding new colors to the spectrum. S.
columnaris has also proved to be an
excellent parent of new cultivars. Again
Karel Lang has used it to great effect
producing, for instance, S. [Prominent
Group] ‘Bohemian Karst’ (2006). This
is one of the purest red colours we have
today. Unfortunately, both parents, S.
columnaris and Saxifraga kotschyi, do
8
not like wet conditions, and neither do
their cultivars. I find it very hard to grow
these plants in our climate. Another
example of what has been achieved is
S. x dinninaris ‘Ray Woodliffe’ (2012),
a cross of S. columnaris and S. dinnikii
raised from wild collected seed.

What lies ahead? I’m sure recent and


future introductions will give us still
more possibilities to create beautiful
new cultivars. Take for instance Tim
Roberts collection Saxifraga TJR 736-01,
9

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probably a hybrid between S. lowndesii better understanding of the Himalayan
and S. alpigena. I think it will make a Porophyllum species.
beautiful parent of new cultivars. Or
Saxifraga ludlowii, another stunning Another puzzle is the nature of Saxifraga
introduction from Nepal. David Walkey hypostoma and the plants that are
already has produced a hybrid using described by Jan Bürgel as Saxifraga
this gorgeous species. It has a distinct x tukuchensis. According to Jan, these
blueish tint. I’m sure that S. ludlowii will plants are natural hybrids between S.
be used for many years to come and hypostoma and S. andersonii. I think
many new spectacular cultivars will be this is not true, based on an article by
introduced. Unfortunately, this species Michio Wakabayashi and Hideaki Ohba
is not readily available yet, so let’s hope (Cytotaxonomic study of the Himalayan
there will be more around in the future. Saxifraga). In this article they show that
S. hypostoma has a different number of
Let us return to our understanding of chromosomes than S. andersonii and
the Porophyllum species. If we look these chromosomes are much larger as
at Saxifraga cinerea for instance, its well. Therefore, it seems highly unlikely
history starts with the description by to me that these two species would
Harry Smith in his book Saxifraga of have hybridised to produce the plants
the Himalaya I Section Kabschia (1958). that were called S. x tukuchensis. We
This description is based on a single definitely need more study to prove the
collection, made by Colonel Lowndes in true nature of these plants.
Nepal in 1950. But what do we know of
the amount of variation in this species One such study was done by a
occurring in the wild? Not much I’m research group led by Natalia Tkach.
afraid. What I do know is that there In December 2015 their article was
is considerable variation among the published in Taxon. The title of this
plants we grow in our gardens today molecular phylogenetic study on
under the name S. cinerea. Often this Saxifrages is ‘Molecular phylogenetics,
variation is explained by assuming that morphology and a revised classification
the plant is not a true S. cinerea but a of the complex genus Saxifraga
hybrid instead. I really don’t know if this (Saxifragaceae)’. In this study over 200
is the actual truth and I believe nobody Saxifrage species were analysed and
does, we simply lack the knowledge to a phylogenetic tree was constructed.
be sure how much variation there is Unfortunately S. hypostoma was not one
in the wild. This is just one example of of the species studied. But it is this kind
how we should be careful about making of study that probably will give us the
judgment on plants from the Himalayas. best shot of solving some of the riddles
It is a pity that so many plants are sold that still exist in our understanding of
without their collection details. We the Porophyllum species.
should all try harder to make sure that
these collection details aren’t lost. We
surely need this information to get a

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