CSR December 2021
CSR December 2021
Contents
6 Click to read 17 Click to read
Arizona, Sansevieria pinguicula
Mexico...Norfolk the ‘walking’
Ian Woolnough Sansevieria
Colin C Walker
Viola coronifera
See Ultra violets Page 51
39 Click to read
Whatever happened to
that Echinocereus?
Peter Berresford
9 Click to read
Orostachys – one, two
or three genera? 21 Click to read
Ray Stephenson Why I don’t like
Lithops
Chris Coombes
51 Click to read
Ultra violets
John Watson and Anita
Flores
28 Click to read
14 Click to read
More diverse
Cumarinia odorata succulents
Graham Evans Jörg Ettelt
Welcome to the December issue of these violas until these were found. Rock Gardener. This covers all 111
the ‘Cactus and Succulent Review’. V. x blaxlandiae (2012) is discussed species, not just those which are
in the article, the other is succulent, and is an eminently
I am delighted to say that I have
V. x zwienenii (2019) shown here. readable and also very important
something really special for you in
work, as it is the first coverage of
this issue.
the subgenus since a one-page
Go to page 51 for the start of ‘Ultra review by Wilhelm Becker in 1925.
violets’, a fascinating article on the It is available to download from the
succulent violas of the high Andes, Scottish Rock Garden Club
written for the CSR by John website.
Watson and Anita Flores.
Finally as we draw to the end of
John and Anita, who are a husband another largely lost year, I would
and wife team, have studied the like to thank everyone who has
subgenus Andinium of the genus contributed in any way to the CSR.
Viola for well over 30 years. During This includes those of you who
that time they have made many have written for me, or supplied
discoveries and are responsible for photographs or drawings,
Viola x zwienenii a natural hybrid
authoring a number of succulent between V. beckeriana and occasionally helped with queries
species including V. lologensis V. atropurpurea. This plant shows and my two hard-working proof
(2011), V. rossowiana (2013), the influence of V. beckeriana. readers.
V. beckeriana (2013), V. anitae (Photo: John Watson)
I would also like to thank all my
(2018), V. abbreviata (2019),
readers for their ongoing support,
V. pachysoma (2019), V. regina John and Anita have recently
without you of course the CSR
(2020) and V. turritella (2020). completed a monograph on the
would not exist.
In addition they discovered and South American Andinopacific
subgenus Andinium of Viola Let’s hope that next year life will
authored three hybrids, two of
published by the Scottish Rock return to normal once more.
which are succulents. Hybrids
were previously unknown among Garden Club with the International Sheila Cude
Back issues
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25 Macleod Road
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www.cactusandsucculentreview.org.uk
4
The cacti of
Cactualdea
Erratum
I would like to apologise for an
error which occurred in the last
issue of the ‘Cactus and Succulent
Review’ September 2021.
Ray Stephenson has pointed out
that in the article entitled
‘Succulents of the Haraz
Mountains’, the plant pictured on
page 43 (shown left) was wrongly
named as Kalanchoe glaucescens.
Kalanchoe fedtschenkoi
In fact it is Cotyledon barbeyi.
Kalanchoe and Cotyledon are
closely related to each other,
however Kalanchoe flowers are
strictly four-partite, i.e. four sepals,
four petals, four carpels and eight
anthers.
Cotyledon have five-partite flowers.
Both Kalanchoe and Cotyledon
have petals fused into a tube.
Photos right: Ray Stephenson Cotyledon eliseae
5
Arizona, Mexico...
Norfolk
by Ian Woolnough
Orostachys –
one, two or three genera?
by Ray Stephenson
Fig. 1
Orostachys
thyrsiflora
resembles the
much larger
O. spinosa and
comes from the
highest parts of
Central Asia
Orostachys – one, two or three genera? continued 10
Fig. 2
Orostachys malacophylla was originally chosen as the type species for Orostachys
Orostachys boehmeri
‘Keiko’ in October
showing the typical
inflorescences
common to the genus.
The herbarium-sheet
displays both plain
green and glaucous
grey plants, so to
distinguish the two
forms horticulturally, I
christened the latter
‘Keiko’ back in 1994.
Fig. 4
Fig. 6
Fig. 9
Hylotelephium cyaneum is similar to Orostachys when out of flower but has little resemblance to say, H. maximum, which is
a tall herb.
Cumarinia
odorata
by Graham Evans
Small, soft, flaccid, difficult, slightly smelly
and seemingly unique
species to Neobesseya, a small genus of diminutive, Escobaria and even possibly Echinomastus or
soft cacti with tubercular furrows, which is often Rapicactus (Turbinicarpus).
considered part of Escobaria today; but again the fruit
The genus Cumarinia was erected by Franz Buxbaum
was different, not the bright fleshy berry of
in 1951, the epithet further highlighting the scent of the
Neobesseya.
fruit. C. odorata became the type species and, to this
The next change came only five years later in 1942 day, remains the only taxon at any rank ever assigned
when Curt Backeberg placed our subject in Neolloydia. to the genus. The combination of its low growing,
Today, Neolloydia is a very clearly circumscribed genus shortly cylindrical, clumping habit, flaccid, long, narrow
consisting of probably two very closely related good and furrowed tubercles ending in a small white areole
species, (and a number of minor variations that have with 7–9 white to pale brown radial spines and 3 or 4
over the years been given species or variety status), much darker, hooked, brittle central spines that are
both of which have fairly large, wide opening purple about 25mm long, small, narrow, dirty pinkish brown
flowers. In Backeberg’s concept, unusually for a man (and, it must be said, fairly insignificant) flowers and
regarded as one of the great ‘splitters’, the genus was small, elongate-cylindric purplish brown and scented
more broadly defined and a bit of a mess including fruits make for a unique and interesting cactus of
species now considered to belong in Coryphantha, broadly mammillarioid appearance.
A mature clump growing at the Wroclaw University Botanical Gardens in Poland (Photo: Agnieszka Kwiecień.
Available under license: CC-BY-SA 4.0)
Cumarinia odorata continued 16
In cultivation, C. odorata is unfortunately not the Although Cumarinia do not have extra-floral nectary
easiest of species to grow and large clumps are rarely glands, the mature fruits are prone to leak and can
seen. I have grown them into 12cm pots but no more, leave small unsightly stains which, if not removed, are
although I did once see an imperfect clump of perhaps also a magnet for fungal infections. All the time the
16–18cm during the dispersal of an old and epidermis is a nice intense green, you should be
established collection. I understand the plant alright, but once a dark, soft blemish appears it is
succumbed shortly afterwards. sadly never long before the whole plant turns to putrid
In my experience, they do best in a very open potting mush. The best policy is to have a smaller plant or two
mix with excellent drainage and in a sunny position growing on to support any older specimen. Whether
with good ventilation. Watering should be light but through nature or poor cultivation, I find the lifespan of
relatively frequent in the growing season; in winter the these plants is no more than 8–10 years.
plants need to be completely dry and as free as Since 1951, the debate has raged tirelessly (or
possible from damp air, but nevertheless they need a tiresomely, depending on your point of view) as to
quite cool rest to ensure dormancy and prevent where this species fits in the classification of the
complete dessication, an amount of which is perfectly cactus family, with various subsequent authorities
normal for C. odorata. It is one of those species for expressing their opinions – and occasionally changing
which it is all too easy to feel sorry, as it shrinks and their minds – based on particular aspects of the plant.
wrinkles in January or February, but any water during Now, at last, it seems that DNA has resolved the
this time will almost certainly prove fatal. It is much situation and preserved the unique position of
better to send it to bed with a goodnight drink, fully Cumarinia odorata (although the possibility remains
inflated, towards the end of the growing season giving that it may be of hybrid origin). n
the pot time to dry out before dormancy.
Growing ghosts
by Michael Madders
Fig.1 Sansevieria
pinguicula in a
27cm diameter pan
Sansevieria pinguicula –
the ‘walking’ Sansevieria
by Colin C. Walker
Fig. 2
purchased larger black-glazed pan. Sadly with sheaths in the manner akin to that of a
this plant is no longer with me since it strawberry plant. The new growth then
succumbed to low or even freezing sends roots into the ground at a raised
temperatures following an unexpected position, thus making the plant look as if it
greenhouse heating failure. The only plants is growing on stilts. As a consequence of
that suffered were half a dozen this, the plant is known as the ‘walking’
sansevierias which should not have been Sansevieria in Kenya.
left in the greenhouse – lesson learned!
My second plant came to me from the
The plant has a very distinctive growth collection of the late Gordon Rowley who
pattern (Fig. 2). It spreads by sending out obtained it in 1993 from the Sansevieria
horizontal runners or stolons not covered expert Juan Chahinian who, in turn, had
Fig. 4
Sansevieria
pinguicula in
flower
Sansevieria pinguicula – the ‘walking’ Sansevieria continued 20
Fig.1
Lithops
verruculosa
‘Rose of Texas’
Fig. 5
Young Lithops
paint brush, encouraging the production of disturbed at a young age. I wait until they
verruculosa ‘Rose
many more seeds. At the BCSS National are at least a year old, but ideally like to of Texas’
Show there were single-headed plants on sow them thinly and leave them to develop
offer, some with colours and patterns that for much longer. Then disaster struck. Jan
even I found attractive. Despite my best discovered the Mesemb Study Group
efforts, many of these somehow smuggled (MSG) and its seed list. (See page 5.)
themselves into the back of the car. So Jan
So she got some shelves. Not just any
stole another area of much needed space
shelves. These were 12 inches wide, 5 feet
from me, and talked about putting up
long and had a total length of 30ft. I was
shelves!
against it at first because of the shade they
The first batch of seedlings germinated so would cast for part of the day over the
well they were soon climbing all over each more worthwhile plants underneath. Plus
other. I resisted the urge to prick them out the fact they would contain Lithops. But
as Lithops do not take kindly to being our garden is south facing so Jan made
Fig. 6
Fig. 7
Fig. 9
Lithops bromfieldii
‘White Nymph’
from MSG seed
Once the first few batches had matured the next MSG seed list and am ashamed to
somewhat I noticed that, despite being the say, ticked a few boxes for myself (Fig. 9).
same species, many of them were subtly
Jan bought more shelves. This time to
different. Some had slight variations in the
span some of the sections on the back and
patterns, others were more distinctly
marked. A few of them were actually quite
attractive (Fig. 7) and I started to have
unholy thoughts of back crossing or
crossing the nicer ones with each other to
encourage further enhancements. The
other advantage of seed sowing was that
very occasionally a weirdo would turn up
(Fig. 8). Meanwhile I stole a sneaky look at
Figs. 10 and 11
Fig. 12
The prizewinning
front of the greenhouse. Same size, but once home, Jan carefully set about colour Lithops display
this time only totalling 15ft in length. I was co-ordinating them with our babies and
abashed. Some of the space was for me. arranging them in a bigger container.
The more mature specimens were now The following year was the BCSS National
potted on into some wonderful square pots Show and the container was looking so
that fitted perfectly on to the shelves and nice that I decided to put an entry in for the
had the extra depth we preferred (Figs. 10 first time ever, not expecting anything but
and 11). just wishing to share the beauty of our
As most of the original plants were now in home grown creations. I can’t begin to
trays or pans anyway, we entered a bowl in describe how elated we were when it came
several local shows and did quite well. second out of a dozen entries (Fig. 12).
Obsession is a strong word but while on
So now I have a confession to make. I have
holiday, why did I spend all my time
to admit that Jan’s little obsession has
searching the beach for rounded pebbles
become a bit of a secret passion of mine
that looked to be the same colours as our
too.
plants? I collected dozens of them and,
Why I don’t like Lithops continued 27
Now one of the highlights of our year is the at the same time. The excitement mounts
MSG seed list. I’m addicted to seeking out when the seedlings finally start to develop
unusual forms such as Lithops julii ‘Hot their patterns and colours. Then we wait for
Lips’ (Fig.13) and Lithops verruculosa the sowing season to come round again...
‘Rose of Texas’ (Fig. 1) and can be seen
But I still don’t like Lithops. And the reason
animatedly pacing along by the shelving at
for that – is because I LOVE them. n
flowering time praying that those I have
selected to cross will cooperate and bloom Photos: Chris Coombes
Fig. 13
Fig. 16
Lithops julii
Fig. 14
Lithops julii
Fig. 17
Lithops lesliei
Fig. 18
Fig. 15
Lithops lesliei var.
Lithops olivacea venteri
A celebration of cacti and succulents 28
Fig. 1
More diverse succulents continued 29
Fig. 5
Bijlia cana
More diverse succulents continued 31
however, are the flower panicles. The La Palma at altitudes of 300 to 800 metres,
succulent petals are a similar mealy-grey to and perennial. The tiny bush should be cut
the whole plant. Touching any part of these back from time to time to ensure compact
plants detracts from their beauty, as the and thus attractive growth; with a bit of
mealy coating can be wiped off. The flower, luck, cut shoots can be re-rooted. The
almost hidden in the bracts, offers a flowers, which appear during spring and
wonderful contrast with its surprising red summer, although not very striking, appear
tones – but you have to look very closely. en masse on the beautiful, compact plant,
They are, like the other species of this and which barely reaches 10cm in height.
other genera, true gems that can be
Keeping members of this genus is a bit
cultivated together with cacti without any
trickier, not only because some species are
problems.
annuals, but also because they need a bit
The Achilles’ heel of the Crassulaceae is more water due to their low succulence,
the spiky, racemose inflorescences, often and should be placed in partial shade. Too
with small flowers, which are not very much water and there is a risk of rotting.
attractive. Such a reduction to the You have to develop a little intuition to be
minimum can be seen, for example, in the able to cultivate these species in the long
genus Monanthes, the species of which are term based on the specific conditions you
endemic to the Canary Islands and the can offer. Most species need a rest in
Selvagens Islands. They are tiny plants with winter and should be watered carefully but
inflorescences that bear only a few flowers, regularly in spring and summer. n
which is also indicated by the genus name, Photos: Jörg Ettelt
‘single flower’.
In the next issue I will look at some beautiful
M. muralis (Fig. 8) is one of the most cacti that give us the pleasure of flowering
attractive members of the genus, native early in the year. After all, anticipation is the
only to the Canary Islands of El Hierro and best of all joys. Monanthes
muralis
Fig. 8
33
March 2021 – no
The house was demolished and the So what I now have, in effect, is a large, house and a view
rubble kept and spread over the footprint open, sunny slope that, with all that
of the house and a couple of metres to the masonry, is going to absolutely bake in
side, leaving me with a southwest facing summer. The high lime content will make
slope where, in places, the rubble is for perfect drainage. To say I am excited by
1.2m deep. the prospects would be an understatement.
May 2021
The sky’s the limit! continued 35
Away from the slope, clearing the we had no idea were present, including a
neglected and massively overgrown site lovely old sundial that had not seen the
has been incredibly hard work but huge fun light of day for decades.
and very rewarding. There were dead trees
Once the site was clear I turned my
and shrubs, self-sown hollies and bay
attention to the planting. I decided to plant
laurel, layered privet, a 12-metre tangle of
mainly, but not exclusively, multiples of
Russian vine that had found its way into
things I know do well here, so I have a
the roof space and five-metre bramble
more limited palette of plants, but used on
stems as thick as my thumb.
a bigger scale, for example a group of five
It was impossible to walk around it at first Agave ovatifolia which should create a
without a struggle. To one side was an large central focus. I had a number of
ancient Buddleja alternifolia that had ‘spare’ potted plants and also, having
spread its wiry branches for four metres in massively overplanted my existing garden,
every direction and had become a huge had loads of plants to transplant. Add to
black hole of light-sucking tangle. We that gifts from friends and I actually had to
disposed of pretty much everything via buy very few new plants. I have a
green waste bins from the council, snipped specimen plant of the slightly tender giant,
or sawn into small bits and rammed in Agave tecta on trial, which will now have to
tight. We have uncovered all kinds of take its chances under the cover of a large
original walls and paths etc, many of which Monterrey cypress.
June 2021
The sky’s the limit! continued 36
October 2021
Some half-dead ‘herbs’ were rescued from also responded beautifully with more
a garden centre compost bin – marjoram, compact growth and more pronounced
thyme, rosemary, hyssop, lavender, curry colour of flesh, tooth and spine and it was
plant and pineapple sage. Immediately the the same with the cacti.
growth shown by these plants was
astonishingly healthy and vigorous. The In amongst the plants I have added some
succulent plants, although not as fast, have drifts of spring bulbs – reticulated iris,
Agave parryi – very black spines developing Agave gentryi – very strong colour especially on the leaf-
bud imprints
Agave salmiana subsp. crassispina – beautifully compact Agave tecta – a potentially massive Agave but a bit tender
and coloured new growth
The sky’s the limit! continued 38
Marcetella moquiniana
39
Fig. 1 Echinocereus
coccineus subsp. coccineus
SB850 Ladrone Mtns.,
Socorro Co., NM JES
96_025
To discuss the current conundrum, it will be necessary extreme east of New Mexico, and south-east into
to focus on the Echinocereus coccineus group (species Texas to Piedras Negras on the Rio Grande. To the
and subspecies) which all fall within Echinocereus east the distribution reached the area north of San
subseries Roseiani. These are characterised by the Antonio and nearly as far north as Abilene. There was
relatively short flower tube, (which can best be also a small extension of the group into the northern
visualised as considering the distance into the flower area of Coahuila and Chihuahua in Mexico (Fig 2).
which a pollinating proboscis has to reach to collect
New discoveries and research have substantially
the nectar).
altered our understanding of this group of plants and
Echinocereus coccineus, in the broad sense, has thick- the distribution of the subspecies. It is probably easier
skinned fruits with a distinct colour change of the to deal with the other members of the Echinocereus
entire fruit at maturity, changing from green to coccineus group before tackling E. coccineus subsp.
yellowish, or pink and orange to red. Echinocereus aggregatus and E. toroweapensis.
subseries Polyacanthi is characterised by a longer
flower tube. The ripening fruits are dark green and, Echinocereus coccineus (Engelmann) subsp.
depending on the exposure to sunlight, either coccineus
completely reddish when ripe or violet with more This subspecies was originally found by Wislizenus
intense sun. during his 1839 wagon train tour from Missouri
I will return to Polyacanthi later. Nobody said that westwards and down through Mexico and described
taxonomy was simple! How much easier life would be by Engelmann in 1848. Plants were found in the pine
if we didn’t have to carry the historic names forward woods of Wolf Creek near Santa Fe, now part of Mora
and used something more meaningful like subser. County, New Mexico.
Breviflora rather than Roseiani and subser. Longiflora Distinguishing features are the number of ribs (8–12),
rather than Polyacanthi! which it shares with E. coccineus subsp. rosei, and the
The known distribution of the Echinocereus coccineus number of radial and central spines (5–12 and
group in 1998 extended from south-west Utah in a 1–normally 3) respectively (Fig. 1). Over the years
wide band down through central Arizona, south of various attributes of the subspecies were recognised,
Colorado Springs in Colorado, covering all but the attracting new names such as melanocanthus,
Fig. 2
cylindricus and conoideus but these are now regarded Echinocereus coccineus subsp.
as synonymous with E. coccineus subsp. coccineus. transpecosensis Blum, Oldach T & J
The altitudinal range is from 1,300 to 2,400m. It is impossible to go any further before noticing the
Echinocereus coccineus subsp. rosei (Wooton ‘elephant in the room’ Echinocereus coccineus subsp.
transpecosensis described in 2015, which affected the
and Standley) Blum & Rutow
mapping of a number of members of what is now
First described as E. rosei by Wooton and Standley in subseries Roseiani. The type location for the plant is
1915, Blum used Kunzmann’s specific epithet as the north of Sierra Blanca, Hudspeth Co., Texas.
base for a new series of Echinocereus, Roseiani. The
holotype was described as being from Dona Ana In 2016 I had the pleasure of staying at a ranch on the
County, New Mexico which borders Mexico just north- banks of the Rio Grande in this county which is
west of El Paso. Clumps may reach up to 30 heads in accessed from Sierra Blanca along a road that soon
size, which is somewhat smaller than 500 which turns to dirt; and the flora becomes distinctly that of
E. coccineus subsp. coccineus may approach but the Chihuahua desert environment. Here Echinocactus
E. coccineus subsp. rosei typically has more central horizonthalonius shares the desert with Dasylirion
spines (4–6) (Fig. 3). E. coccineus subsp. rosei also wheeleri, Escobaria tuberculosa and several
favours lower altitudes from 1,300 to 1,600m. The echinocerei including E. enneacanthus, E. stramineus,
mapping of Echinocereus coccineus subsp. coccineus E. dasyacanthus and E. coccineus subsp.
and rosei as a ‘group’ has changed little since 1998 transpecosensis. Revisiting in late April 2019, many of
but the known area of subsp. coccineus has risen to the cacti were in flower (Fig. 4).
greater prominence at the expense of subsp. rosei, By the ranch along the river there were some different-
once thought to extend further north. looking plants which we believed to be a transitional
Fig. 3
Echinocereus coccineus subsp. rosei with Swallowtail butterfly pollinator. Franklin Mountains State Park,
El Paso Co. TX. 1,523m
Whatever happened to that Echinocereus? continued 42
Fig. 4
Echinocereus coccineus subsp. transpecosensis, west of Sierra Blanca, Hudspeth Co. 1,341m
form between E. coccineus subsp. transpecosensis subsp. transpecosensis lies between the two previously
and subsp. rosei (Fig. 5). described subspecies above and the two which follow.
This subspecies is described as having 7–11 ribs, 6–11 All the field-work and study undertaken since 1998 has
radial spines and 1–4 centrals. I am in some doubt as rendered the previous name E. coccineus subsp.
to the criteria regarding central spines here. In 2008, I aggregatus redundant and has also had an impact on
first saw plants in the Sierra del Carmen in Coahuila, the mapping of the following two subspecies (Fig. 7).
subsequently revisiting them on several occasions.
Here E. coccineus subsp. transpecosensis frequently
displays 0–1 central spines, although the rib count is
within the recently documented criteria. In those days
we were calling this plant E. coccineus subsp.
aggregatus (Fig. 6).
In 2017 Blum et al. reported on the results of a new
characteristic that helps to differentiate the subspecies
of the ‘claret cups’. The spine surface was found to be
consistently different between certain subspecies,
which proved critical in understanding the relationship
of plants in this subseries. Spines from subsp.
transpecosensis have a smooth-tubular, grooved
epidermis with an incipient tuberculate structure (the
beginnings of small, rounded projections). E. coccineus
subsp. coccineus and subsp. rosei have, by contrast, a
Fig. 5
smooth, striated spine surface whereas E. coccineus
subsp. paucispinus and subsp. roemeri both have a Transitional form of E. coccineus subsp. rosei and
spine surface which is significantly tuberculate. Both E. coccineus subsp. transpecosensis,
geographically and developmentally it would seem that Hudspeth Co. TX. 1,152m
Whatever happened to that Echinocereus? continued 43
Fig. 6
Echinocereus coccineus subsp. transpecosensis in the Sierre del Carmen, Coahuila displaying 0–1 central spines
Fig. 7
Fig. 8
Echinocereus coccineus subsp. paucispinus, Evans Creek off US90, Val Verde Co. Texas
Echinocereus coccineus subsp. paucispinus Grande, with Jos Huizer and was fortunate to find
(Engelmann) Blum, Lange & Rutow some close to the type locality where the Pecos River
Cereus paucispinus is one of Engelmann’s finds spills into the Rio Grande. If you blink, travelling down
described in 1857 but the combination with US 90, it is easy to miss these plants – far better to
Echinocereus coccineus is attributable to Blum et al. in turn off when you reach the area and search slowly,
1998. The mapped distribution of this subspecies has then you will find them under rock overhangs (Fig. 8)
reduced by some 75% since the combination was between 250 and 500m elevation.
created. Plants growing to the south and east of the This is possibly the easiest of the subspecies to
1998 distribution are now attributed to E. coccineus identify, having very few ribs (5–7) and 0–1 central
subsp. transpecosensis. Over 20 years ago, I went spines accompanied by 3–7 radial spines. The specific
looking for these as I travelled south, following the Rio epithet translates as ‘few spines.’
Echinocereus
coccineus subsp.
roemeri
(Muehlenpfordt) Blum,
Lange & Rutow
Originally described as a
Cereus in 1848, a
combination of this name
was never made with
E. triglochidiatus so the
taxon has made fewer
‘hops’ than many of its
relatives. Like subsp.
paucispinus its range is
quite low (between 300
and 600m) and it relishes
Fig. 9
Echinocereus coccineus
subsp. roemeri accompanied
by E. reichenbachii subsp.
caespitosus Medina Co.
Texas at 351m
Whatever happened to that Echinocereus? continued 45
Fig. 10
either granite or limestone rocks. The western edge of Plateau. It is easier to distinguish from subsp.
its distribution has been ‘eroded’ by the relatively new paucispinus as all the key criteria are larger (rib count
subsp. transpecosensis but it grows in the area north ranges from 7–10, radial spines from 6–9 and centrals
of San Antonio on the more temperate Edwards from 1–4). There is no need to pack litres of water
Fig. 11
Whatever happened to that Echinocereus? continued 46
Fig. 12
when you go into the field here, the meadows and interest are the longer flowers which can be
roadsides are full of annual and perennial flowers hermaphrodite or functioning female or male flowers
(Fig. 10), most of which do not originate from succulent (Fig. 11). Plants possess 9–11 ribs, 7–11 radial spines
plants. It is not difficult to find these cacti where rock and a normal compliment of 1–2 central spines,
outcrops occur by the side of smaller farm roads. They although exceptionally this may be as many as four.
are often found growing in granite or limestone in
tandem with E. reichenbachii subsp. caespitosus
between 300 and 600m (Fig. 9).
To explore what happened to Echinocereus
toroweapensis which, according to the Echinocereus
1998 monograph, covered most of central and north-
western Arizona (Fig. 2), a good understanding of the
second subseries of series Roseiani, namely
Polyacanthi, (with its accompanying longer flower
tubes) is required. Back in 2003 I explored a large area
of Arizona and confidently labelled all the large clumps
of red-flowered plants in northern Arizona as
E. toroweapensis. The revelation of Echinocereus
bakeri, first described in 2015, had a profound effect
on the understanding of the red-flowering echinocerei
in central/northern Arizona.
Echinocereus bakeri Blum, Oldach T & J
The plant identified and collected by Marc Baker was Fig. 13
named in honour of his extensive contributions to a
Echinocereus bakeri Barnhardt Trail, Gila Co. Arizona
better understanding of the genus. Of particular 1,251m
Whatever happened to that Echinocereus? continued 47
Fig. 14
Echinocereus canyonensis, Toroweap Point, North rim of Grand Canyon, Mojave Co., Arizona. Shown by kind permission of
Traute Oldach
It is rare to see one or two stems of this plant but In 2003, during travels through northern Arizona, I
clumps of more than 30 heads are very common struggled to differentiate E. triglochidiatus subsp.
(Fig. 13), and plants can reach 500 stems. It is no mojavensis from E. toroweapensis but eventually
surprise, coming from this part of the USA, that started to recognise the subtle differences, particularly
E. bakeri grows at altitudes between 500 and 2,450m the different spacing between stems in a clump.
(Fig. 12). The spine surface is also indicative of Clearly, I had not read Blum’s 1999 article before my
separation as a species from previously discussed 2003 visit, which identified E. toroweapensis as a
taxa, having distinct longitudinal grooves with the synonym of E. canyonensis! The type locality was only
beginning of small tuberculate growths. 40km apart for the two names. In the circumstances,
the earlier name of E. canyonensis took precedence
Echinocereus canyonensis Clover & Jotter
and E. toroweapensis ‘disappeared’. Imagine how
The presence of another red-flowering Echinocereus many plants I would find in 2003 without a name to
belonging to subseries Polyacanthi, inside the differentiate them!
distribution of E. bakeri (Fig. 7), is most interesting and
there is a connection here with our other ‘disappearing’ By 2017, after a lot of further work, the small area of
name, E. toroweapensis. E. canyonensis (Fig.14) was distribution of E. canyonensis was mapped as being
described in 1941 having been collected on steep within the distribution of E. bakeri and well-
limestone outcrops ‘100 yards from the river at Bass differentiated by several factors:
Cable below Hermit Creek Rapids in Grand Canyon, E canyonensis E bakeri
Coconino County’. In Blum et al (1998), this plant was Ribs 11–14 9–11
relegated to potential synonymy with Radial spines 9–13 7–11
E. triglochidiatus subsp. mojavensis, another Central spines 4–7 1–2 rarely –3 or 4
widespread ‘claret cup’ ranging from central California, Flower length 30–45mm 50–70mm
through Nevada, Utah and as far east as western Flower diameter 15–35mm 35–45mm
Colorado and down into northern Arizona. Nectar chamber 2.5–6mm length 6–9mm length
Whatever happened to that Echinocereus? continued 48
Fig. 15
Echinocereus yavapaiensis M.A. Baker penetrating gaps between boulders or in rock faults
(Fig. 15).
In 2006 another ‘claret cup’ was described from
Yavapai County, the Arizonan county above Phoenix. The dichotomous key produced in Blum et al (2017)
This is relatively easy to reach and by 2013 was on my indicates the defining characteristics of this species to
schedule to visit. This is another long-tubed flowering be its ploidy and the larger size of its flowers
Echinocereus within subseries Polyacanthi. (35–80mm long x 35–50mm diameter). The rib count is
As with E. canyonensis, the distribution area from 9–14. Radial spines number 7–10 and central
(Fig. 7) falls within the much larger range of E. bakeri. spines between 1–6 (Fig. 16).
One should exercise caution before labelling
photographs in northern Arizona! Unfortunately, the plants had long since finished
flowering by the time of my visit towards the end of
Unlike the other plants in this article which are all May but plump fruits had formed on several plants
tetraploid, E. yavapaiensis is hexaploid. Both functional (Fig. 17) and these proved to contain some ripe seeds,
male and female flowers are produced as well as although not as many as one might expect when
hermaphrodite flowers. From observations north of dissecting the fruit of a ‘claret cup’. The seed was left
Phoenix, it is clear that the plant has a strong at appropriate-looking sites for germination and
preference for volcanic rock and grows with its roots development at the location.
Whatever happened to that Echinocereus? continued 49
Fig. 16 Fig. 17
Clump of Echinocereus yavapaiensis north of Phoenix Echinocereus yavapaiensis fruits May 2013
Echinocereus santaritensis W. Blum & Rutow The first time I saw it in 1997 was completely
This is the antepenultimate species, again belonging to unplanned. I was following a lead on Echinocereus
subseries Polyacanthi and the most southerly within bonkerae which was reported as growing along the
Arizona, stretching over the border down into Sonora Salt River on White Mountain Apache land in Gila
and Chihuahua. The original description of 1998 had to County. It was late March and both plants were in
be corrected (Blum 1999) as the herbarium reference flower, although E. santaritensis was an unidentified
was incorrectly cited for the holotype. red-flowering Echinocereus at the time (Fig. 18). The
republished description for the plant in Blum et al
It is named after its occurrence in the Santa Rita (2017) indicates it flowers from April to May, so these
Mountains from where the holotype was collected. flowers were early. It is the long flowers and the fact
Fig. 18
Ultra violets
by John and Anita (Flores) Watson
An introduction to the succulent violas of the high Andes
‘Viologists’ is what our late lamented colleague Kim Blaxland called herself
and the two of us. The genus Viola, or one part of it, a South American
Andinopacific group, has been our principal focus of field exploration and
investigation for well over 30 years.
For obvious reasons we call them nectar-bearing spurs extending back from
‘sempervivoids’, and they attain their the lowermost petal. V. sempervivum has
maximum physical expression in our ‘one smallish yellow flowers, those of
under two dozen’ described below. These V. santiagonensis being larger and white.
range along the southern half of the overall
The remaining species, V. portulacacea, is
distribution of subgenus Andinium, from
flat to the ground and, although not
central Chile and Argentina southwards,
preserved in flower, was reported to have
where one of them, V. auricolor, a low, mat-
pale blue corollas. We have searched for it
forming species with showy bright, pale
thoroughly for a long time, exactly where it
orange corollas, is the most southerly of
was collected, but in vain. The valley is
the subgenus.
now shrub-covered and we believe it was
Many species of the subgenus are swept down by a land slip from higher
exceedingly rare, including 20 only ever ground above, either as plants or seeds.
found once and not seen since. Some of V. lologensis, another considered to be
those are now probably extinct. Four extinct and which we have also searched
belong to our group. One, V. comberi from for fruitlessly, was found in the mid-1920s
Argentinian northern Patagonia, resembles by the British horticultural plant hunter,
golden-yellow flowered V. coronifera and in Harold Comber, as two closely adjacent
fact shares (or shared) the same level colonies in Argentinian Patagonia.
mountain top. They can only be told apart
The rosulate violas under consideration
by close examination of herbarium
come in two forms, either with their
specimens.
rosettes short or even flat to the ground as
The others are from the Santiago in V. abbreviata, V. auricolor, V. coronifera,
mountains of central Chile. Two have long V. cotyledon, V. dasyphylla, and V. regina,
Volcan Copahue, location of V. pachysoma, V. cotyledon and their hybrid V. x blaxlandiae (Photo: John Watson)
Lago Vintter, Argentina, location of V. columnaris and one of the southernmost locations of these violas (Photo: Harry Jans)
Ultra violets continued 54
Evolution
So why and how did these particular violas
get to be so different from others of their
kind, and what is their evolutionary origin?
Thanks to modern molecular analysis we
know where, when and under what
circumstances this happened, which
enables us to provide speculative but
realistic answers to those questions.
The origin of the Violaceae has been
determined to have taken place in what is
now the south of South America. At the
time it was covered in tropical forest and,
as now, except for most of Viola, genera of
the Violaceae family consisted of short
shrubs or subshrubs. We may reasonably
assume they grew in any habitats where
sufficient light was available, such as
clearings and by riversides. It was from
among these that the first violas, also more
or less woody, evolved.
Then, around 34 million years ago, a
dramatic global cooling event gave rise to
Antarctica, converting the rain forest of the
southern half of the subcontinent to plains
and local sectors of temperate woodland.
During the same period, tectonic collision
along the Pacific coast of South America
resulted in the initiation of precipitous
localised vulcanism and the gradual
orogeny that gave rise to the Andean chain.
Inevitably, most old habitats were
destroyed and new ones created. Only
species which could adapt to the abrupt
change and those newly evolved to take
advantage of it survived.
Viola proved to be particularly adept. As
well as retaining elements of its woody
features to this day, a new line evolved
which was able to occupy the chemically
rich but hostile and otherwise almost
vacant ash and pumice fields of the new
volcanoes, most or all of which were still
active at the time. They were the ancestors
of our succulent sempervivoid species; in
fact the very first herbaceous violas.
Violets or pansies only came into existence
millions of years later, so our sempervivoids
must have evolved from the woody
conglomerate, although we have no
present chain of intermediate forms to
confirm this. Continuing frequent and
violent eruptions certainly extinguished herbaceous violas reached the lands of the
large numbers as well as providing new northern hemisphere and spread rapidly to
habitats, which led to further evolution, and cover almost every part of it except
among those eradicated were all but one of deserts, tropical forest and the Arctic.
these ‘intermediates’. That species,
Meanwhile, our particular subgenus has
V. fluehmannii, a dwarf, ericoid shrublet,
gone on evolving to produce many new
exists to the present, mainly on some
and varied forms, including annuals, these
Araucaria (monkey puzzle) clad volcanoes
latter reaching down as far as the Pacific
of near-southern Chile, but also in adjacent
coast. Our succulents, however, are only
Argentinian north Patagonia.
found sparsely to the north of mid-Chile
Harvey Ballard of Ohio University and his and adjacent Argentina as a mere five
colleagues have shown that ancestors of species up to the equator. In all probability
the Hawaiian violas were transported there they were continuous once, but the
across the vast Pacific reaches by birds. extreme uprise of the central Andes (the Viola fluehmannii
(Photo: John
Without doubt, our subgenus Andinium Altiplano) simply proved to hostile for them,
Watson)
species were, and probably still are, being as for almost all others of their kind.
spread along the Andes, predominantly
from south to north, evolving as they went,
to meet the new geoclimatic conditions of
Andean uplift. Whether they reached the
top limit of the subcontinent and were
wiped out above Ecuador by the Gulf of
Mexico region meteorite effect, or were
unable to inhabit the Andes above the
equator, we do not know. For certain
though, about 16 million years ago
V. fluehmannii often grows in open groves of monkey puzzles (Araucaria) on southern Chilean volcanoes.
(Photo: Anita Flores)
Ultra violets continued 56
Viola dasyphylla (Photo: Anita Flores) Viola aizoon (Photo: Anita Flores)
Viola leyboldiana (Photo: Anita Flores) Viola petraea (Photo: Anita Flores)
Viola beckeriana (Photo: John Watson) Viola skottsbergiana (Photo: Anita Flores)
Ultra violets continued 57
Viola coronifera
Anita exploring an
Poor little blighter. Argentinian
It’s suffering as Patagonia
much as we were. V. coronifera
But at least we habitat in
could go back to a unseasonally
warm hotel room wintery late spring
for the night! weather. (Photo:
(Photo: John Watson) John Watson)
Viola coronifera
(Photo John
Watson)
See also the front
cover
Ultra violets continued 58
Lago Teno,
an atypical high
Andean habitat of
V. cotyledon in central
Chile. It is usually
found lower down in
Patagonia
(Photo: Anita Flores)
Viola cotyledon
(Photo: Anita
Flores)
Environment
Given this picture of their evolution, history
and distribution we can place their
Several succulent
particular succulent, rosulate form into a
species of the Bromeliaceae, notably
particular environmental context. Most
puyas, also inhabit semi-arid sectors in the
succulent plants have evolved to cope with
same regions. These places are not
heat and prolonged periods of drought,
populated by violas, however, and climatic
storing water in their bodies (stems) or
aridity is not the reason for their
leaves, or both, until the next precipitation
succulence.
eventually arrives. As readers will obviously
know, South America is a major centre of Volcanic ash and pumice drain very rapidly
distribution for the cactus family, which and soon dry out superficially: but our
even inhabits the Atacama Desert there. violas have deep vertical roots which reach
Ultra violets continued 59
The upper Andean site, Los Andes province, Chile where our friend Carlos found V. regina.
Inset: Viola regina. (Photos: Carlos Celedón)
down to the constantly moisture-laden levels of the Andes are extreme, with
levels up to a foot or more below the summer heat, snow in winter and powerful
surface which supplies them with all they winds. No typical herbaceous plant can A small colony of
need. But climatic conditions on the upper survive these, for which reason these high V. rossowiana in its
points are largely uninhabited. The very extensive type
sempervivoids, however, have evolved the site, extreme
dense, ground-hugging, succulent rosettes northern Patagonia,
to give them the resilience to combat these Argentina
conditions. They are immune to any wind (Photo: John Watson)
force and their sturdy form protects them Inset:
Viola rossowiana
(Photo: Anita Flores)
Ultra violets continued 60
from all the conditions Mother Nature ground-covering herbage, and for this
throws at them, except very occasional reason they are usually found in splendid
landslides and volcanic eruptions. Even the isolation or in bare stretches between
last named only rarely destroy more than a patchy dwarf high-altitude central Andean
part of a local habitat. The need for the or Patagonian vegetation. It also explains
maximum of exposed leaf area to enable why they only occasionally occur lower
photosynthesis is also paramount, and for down than mountain tops.
this the overlapping rosette is
Exceptions to that generality are the narrow
unsurpassable. Their form to suit their
leaved species V. dasyphylla,
environment could not be bettered.
V. aizoon and V. cotyledon, which can
One aspect of the alliance is their inability tolerate a more significant neighbouring
to tolerate competition from vigorous flora. Their more attenuated leaf-shape and
Viola abbreviata
The same plant with
our jeep keys to give
an indication of its
remarkably tiny size
(Photo: John Watson)
Ultra violets continued 61
Our jeep churning up dust on our way to be shown a new viola which we described as Viola turritella. (Photo: John Watson)
Viola turritella
(Photo: John Watson)
Ultra violets continued 62
Viola anitae (Photo: John Watson) Viola micrantha, the only succulent annual
(Photo: David Haselgrove)
Viola polycephala,
Similar in form but not rhizomatous is Northernmost of all is Ecuadorian endemic the northernmost
V. sacculus with showy white flowers. Both V. polycephala with its remarkable blackish of all the
inhabit Patagonia. V. micrantha, widespread green and yellow flowers. The numbers in rosulates, almost
but intermittent from extreme north-west the tropical Andes are completed by reaches the
Argentina to central Peru, is the only V. bangii, V. pygmaea and V. pusillima. equator
succulent annual. (Photo: anon)