Cistanthe Celedoniana.

International Rock Gardener
ISSN 2053-7557
Number 132 The Scottish Rock Garden Club December 2020

---International Rock Gardener---
December 2020
I doubt that many people will have reason to look back on this year with any
pleasure – as the ‘Year of the Covid19 Pandemic’ there has been too much loss
to engender fondness in most hearts. Family members and friends have been
taken by the disease, disruptions of all kinds have ruined plans for events, travel
and projects around the world in every sphere. Even in the year to come, it is
unsure if the way of life for all, including plants lovers will be able to proceed in
any form that we have come to regard as ‘usual’ – instead we must continue to
find ways and means to discover some form of ‘new normal’ – not the happiest
of prospects but there have been great strides made with international internet
meetings which could remain as we get to grips with new possibilities to make our existence bearable.
I do believe that those with an interest in plants and the natural world have an advantage in having
something so hopeful in these times. Our increased concentration and study of our plants over the
recent lockdowns has meant many are understanding the needs of the flora and fauna around us as
never before – there also seems to have been a positive explosion in the level of interest in
gardening and self-sufficiency over the last few months. How fortunate those of us with our own
gardens really are - what a pity it is has taken a pandemic to highlight that! M.Y.
Cover photo: Argylia adscendens var. adscendens Christian von Bohlen
This month’s IRG presents an article by John and Anita Flores Watson on Cistanthe celedoniana
(Montiaceae) which is a new species from Valparaiso Region, Chile, with notes on the genus, including
a new combination, and photos of the flora leading up to the type area in the upper Río Aconcagua
Valley. It is a pleasure to have Anita and John
able to write again after their brush with Covid-19.
Our second submission this month is from
Panayoti Kelaidis, one of the most famous of
American horticulturists, about Alan McMurtrie,
christened by Panayoti as the King of Reticulates!
Since 1980 Panayoti has represented Denver
Botanic Gardens in multifarious endeavours as an
expert in horticulture. His many other talents
include writing and numerous awards, such as
two of the highest honours in American
horticulture. Praise from PK, as he affectionately
known, is therefore praise indeed!
Panayoti Kelaidis in China
Alan McMurtrie is one of the world's
foremost breeders of reticulata Irises. He
has received awards for his work from the
likes of British Iris Society and his hybrids
excite interest wherever they are seen. Alan
has travelled widely to give talks and seek
commercial growers for his hybrids. Alan’s
website is at http://www.reticulatas.com
Alan McMurtrie Iris ‘Storm’
- and good health in 2021!

www.srgc.net Charity registered in Scotland SC000942 ISSN 2053-7557
--- Species Description ---
Cistanthe celedoniana (Montiaceae), a new species from Valparaiso Region,
Chile, with notes on the genus, including a new combination, and an illustrated
account of the flora leading up to the type locality in the upper Río Aconcagua
Valley.
John Michael Watson1*, Arve Elvebakk2, Christian von Bohlen3 and Ana Rosa Flores1.
1
Casilla 161, Los Andes, Región de Valparaiso, Chile:
e-mail ⁓ john.anita.watson@gmail.com.
2
University of Tromsø - Arctic University of Norway, the Arctic University Museum of
Norway, P.O. Box 6050, N-9037, Tromsø, Norway:
e-mail ⁓ arve.elvebakk@uit.no ORCiD id: https://orcid.org/0000-0002-7682-3797.
3
San Fransisco J42, Hijuelas, Región de Valparaiso, Chile: e-mail ⁓ chvonbohlen @gmail.com.
* author for correspondence.
Published in International Rock Gardener (IRG) 132, December 2020, pages 3-50.
Summary
We begin by profiling the late Carlos Celedón, who

lived close to John and Anita, and who discovered
the type site of Cistanthe celedoniana, the species
presented here - which is named in his memory. It is
the last of two species to be described for botanical
science as discovered by him while exploring on his
own. However, his achievements went well beyond
that, He encountered a 'long lost' annual rosulate
viola at two sites in the general neighbourhood, and
took John and Anita to one of them to identify it for
him. While there they recognised three other
species in the unique restricted habitat as new to
science. It is for these outstanding achievements as
a keen amateur naturalist and explorer, and also in
memoriam for his lamented early death that we
name the novelty Cistanthe celedoniana for him.
None of the present authors have seen the plant at
the type site, although John and Anita have grown
containerised specimens in their garden. However,
they, the Watsons, have travelled along its very
narrow approach road on several occasions,
originally to take Carlos there, and have
encountered and photographed wayside plants that
flower at the same time as the cistanthe. These and
their habitats are narrated in the main part of the
introduction and background.
fig.1: Carlos Celedón, the main protagonist, who

discovered the species named for him herein.
(30 Nov 2014. JMW).

Very recently a second population was reported by friend, colleague and fellow author here,
Christian. It's sited some 70 km to the SSW of the one found by Carlos.
We cover all the formalities of the new plant in the second botanical part, beginning with Taxonomy.
Here, friend and colleague, Norwegian academic botanist Arve, joins in. He has made an in-depth
study of the cistanthe's systematic context, which is also included here. In this second part we also
elaborate on two interesting phenomena observed at the type site: pollination of the flowers by ants,
and diurnal movements of the leaves and flower stems, although this last has also been noted by
Christian at his location.
Introduction and background
The most important person by a long chalk in this account is the late Carlos Celedón of Los Andes
[figs.1, 76, 83]. A fanatical amateur naturalist and fine photographer, he discovered the type site of C.
celedoniana during his regular explorations in the general area within a roughly 15 km radius from his
home, which ranged between due south to due west towards the border with Argentina. It included
the high lateral cordillera to 3000 m-plus bordering the upper Aconcagua valley to the south. Regular
readers of the IRG have already been introduced to Carlos twice, in 2018 and 2020, and he has
appeared earlier in the SRGC journal, in 2014. See the following paragraph for full references of
these.
He and the Watsons first met in the second half of 2013, precisely in the context of the last-cited
exploit. Carlos was an intrepid and tireless hiker and climber who would walk a fair distance to the
foot of an Andean peak, climb to its summit ridge some 2250 m above him, then return back down
and home, all in a day, including much photography. In that way he found his exciting new rosulate
Viola regina [figs.80, 81] (Watson & Flores 2020). But it was to another annual species much lower
down, V. chamaedrys [fig.78], that he took the Watsons on that first occasion (Watson & Flores
2014). In fact it had not been seen again since shortly after its initial discovery 150 years ago - so a
valuable revelation by him. He had first noticed it when young with his father looking for indigenous
artefacts, and now wished to know its identity, so he asked a botanical friend who also happened to
know John and Anita. He was told he lived next to those who knew more about those violas than
anyone else - the Watsons! When they got to know the location, a remarkable close-set group of
unique bare clay habitats, they discovered three new species, one the delightful dwarf Alstroemeria
piperata [fig.79] (Watson and Flores 2018), the other pair still awaiting publication. It's partly in
recognition of this significant legacy for botanical science that the new species bears his name.
But the original intention for the specific epithet was C. gibbosifolia, referring to the unusual and
almost unique leaf formation for a cistanthe. John and Anita encountered the shock news of his death
by e-mail while in England visiting their daughters. Apart from being phenomenally active, he was
only 46. But unfortunately he'd chain smoked from his teens onwards. Enough said. The Watsons
had been looking forward to years to come of cooperation, with him as their legs and eyes and them
as his botanists. But alas, it was not to be. The best laid plans ... (an appropriate quote for Scottish
Rock!).
The species is endemic not only to Chile [fig.2], but also to that country's Valparaiso Region [fig.3].
Type locality ‗A‘ is situated 25 km due east from Los Andes towards the Chile/Argentina border in the
upper Aconcagua River valley, which is hemmed in on both sides by two parallel lateral Andean
ranges [fig.4]. The other known location of C. celedonia lies well to the SSW – ‗B‘ in the Reserva
Ecológica Oasis de La Campana [figs.3, 4]. The two locations are remarkably far from one another,
the approximate distance between them being: 70 km [fig.4]. This indicates with little doubt that there
must be further undiscovered populations connecting them in unexplored parts of the E-W foothills,
probably a significant number.

fig.2: Southern South America, Chile. Valparaiso Region - where
Cistanthe celedoniana (circled cross) is endemic - arrowed and
marked red.
fig.3: Valparaiso Region and the two distinct known locations of

Cistanthe celedoniana there, A, the type, and B, La Campana.
(Courtesy of Touristel, Chile)
fig.4: Type location of Cistanthe celedoniana as asterisked green circle and Christian's as asterisked
orange circle. (Courtesy of Google Earth)
fig.5: The lower Río Colorado seen from the approach road to the river crossing, with the cistanthe
type site beyond. (3 Nov 2020. JMW)

Following the little red river
Natural water courses in Chile which become reddish brown with mud, at least when in full spate, are
often called Río Colorado. One such, a large tributary draining down from the Santiago Andes is very
well known. But there is a picturesque much smaller Río Colorado closed in by steepish valley sides
which flows swiftly from north to south down through the foothills [fig.5] and joins the main Río
Aconcagua 25 km to the east of Los Andes and higher up. In fact it's so small it might get called a
stream in Britain. On its west bank runs a narrow dead-end adopted road, just about wide enough for
two reasonably sized vehicles to pass in most places. It ends its availability as a public thoroughfare
after just over two kilometres, when it continues on the east bank as a private unmade access track
to a small mining operation [fig.17]. Occasional modest country dwellings are scattered along the way
on both sides of the tarmacked stretch. Trees clothe the immediate surrounds of the water in most
places, and - if cars can be left in roadside clearings above - it's a favourite picnic spot where people
scramble down to level shady spots by the waterside. C. celedoniana is reached by following the
public road to its limit, obtaining permission to cross the bridge over to the shrub-clad eastern side,
walking or driving along the private track for about two and a half kilometres to where the lower, type,
population was found in 2012. [figs.19, 20].
Carlos didn't drive, his family had no car, so he used to hitch-hike to his footslog starting points, in
this case the small settlement of Río Colorado at the confluence outlet of the tributary of that name,
25 km east of his home. By good fortune he managed to get consent from the owner of the house by
the bridge and she opened the gate for him to carry on up along the private track. Although a much
shorter sortie and a more level one by far than was his usual wont, it was one of three he visited to
yield species new to science.
The only one of us four to have seen C. celedoniana in situ is Christian. Nevertheless, John and
Anita have travelled up its Río Colorado access road on a number of occasions, including as recently
as early November of this year (2020). On several occasions they have carefully passed and greeted
traditionally clad and friendly huasos on horseback [fig.6]. They have also carefully noted and
photographed all the flora as depicted and described herein, where the taxa are arranged from the
first encountered
near the start to that
seen nearest the
public thoroughfare's
dead end.
fig.6: A friendly
huaso (traditionally
dressed Chilean
'cowboy') met along
the Río Colorado
road.
(30 Aug 2020.JMW).

Alstromeria pulchra subsp. pulchra [fig.7] was the
first to be seen along the way, but appeared here
and there for almost the full extent of the road. Over
and above its register in five central regions, making
it one of the commonest of the genus in Chile, it has
excelled itself this year, appearing in astonishing
quantities, especially along the coast. The low, white
flowered composite shrub Baccharis salicifolia
[fig.8], with its bunched inflorescences like tiny
untidy brushes, lined the roadside towards the river
abundantly here and there, and its late flowering
attracted many butterflies of at least three species.
One of these was the fritillary Yrameae cytheris,
[fig.8], whose caterpillars, like all those of its small
genus, feed exclusively on violas. It therefore
informed the Watsons that those plants, their main
focus of study, must be nearby - but none were seen
... then.
fig.7: Alstroemeria pulchra, scattered along the

higher, more exposed W wayside of the river valley.
(31 Oct 2020. ARF)
fig.8: Yramea cytheris, the southern fritillary

butterfly, on Baccharis salicifolia. The fritillary larva
feeds on violas. (3 Nov 2020. JMW)
Attractive, familiar Calceolaria polifolia [fig.9], a

neat, dwarfish shrublet with whitish grey leaves
covered densely in short hair, sought higher, drier,
shallow rocky inclines - exposed and with little or
no competing flora. Another identically sized and
equally compact shrublet with pink flowers
occupied the same habitat, but the two were never
seen together. That second species belongs to a
small family endemic to South America below the
equator, one which probably nobody who grows
plants will have heard of - the Francoaceae. It's
also a member of an equally small genus,
Viviania. Curiously, flowers of these fairly similar
pink or white species have five petals, dry, even
when fresh, so if they are picked and not put in
water, they will still make an everlasting bunch in a
small vase. Their main drawback, as can be seen
from the photos here, is their untidy-looking
inflorescences. The one in the Río Colorado valley
is V. marifolia [figs.10, 11].

fig.9: Calceolaria polifolia, one of Chile's few small shrubby species, on the upper, exposed roadside
bank. (3 Nov 2020. JMW)
fig.10: Another small shrublet, Viviania marifolia, in the identical Calceolaria habitat.
(1 Nov 2020. JMW)
fig.11: A close-up of the papery flowers of Viviania marifolia. (7 Nov 2020. ARF)
fig.12: Haploppapus velutinus is fairly common at mid-

elevations in the upper Aconcagua valley.
(7 Sep 2020. JMW)
We're now getting close to the point where the public

road ends and continues as a private track across the
other side of the river via the small bridge. In that
ultimate stretch three petaloid species and two
pteridophytes were seen. The first of the flowery trio,
Haplopappus velutinus [fig.12], a composite, hadn't
been seen anywhere before by John and Anita,
although it has turned up this season almost
everywhere they explored in their sector. However,
Haplopappus are umpteen and legion and not easy to
identify unless one deliberately spends time doing so.
This species was a brute to photograph. It produces
flower heads in a long succession. The first to appear
are few and scattered. When enough have opened to
fill a frame effectively those earlier ones have shrunk
to unsightly brown, shrivelled blobs, so the only
recourse is to 'dead head' the plant thoroughly,
otherwise it's a case of just photographing the natural
mess!

fig.13: The well-known Tropaeolum tricolor, here in unusual prostrate mode. (30 Aug 2020. JMW)
fig.14: Although recorded from 7 regions up to 2600m,

Adesmia mucronata doesn't appear to be common.
(30 Aug 2020. JMW)
Familiar, common and hummingbird-adapted

Tropaeolum tricolor [fig.13], with little doubt the most
widespread of its genus, has a north-south range of
some 2350 km and occurs laterally from near sea
level to 2800 m in the Andes, yet remarkably is
confined within Chile! It's particularly abundant in the
upper Aconcagua Valley sector, where it may be seen
draping shrubs and fences by the wayside. Often T.
azureum grows mixed in with it, but that plant is one
of three of their subsection Chilensia not to have
produced a hybrid, the others being Tropaeolum
austropurpureum and T. kingii. The latter has never
been seen with another tropaeolum anywhere near,
so hybridization is out of the question. The reason for
the failure of the other two to cross with adjacent
species is straightforward and obvious. Their
markedly different blue-range colour from the rest
(which are yellow or red) attracts distinct pollinators,
various bees, which do not visit the other colour range
any more than the hummingbirds visit the blue or
purple flowers. Both species are established in
cultivation, and we hope one day someone may try
crossing them artificially. T. tricolor at Río Colorado was interesting to record in that it frequently
trailed over the ground, something rarely seen elsewhere. The last species encountered in bloom
was a very attractive, erect, herbaceous Adesmia with an open spike of typical yellow pea blossoms.
Again it was new for the Watsons, and after considerable research they managed to identify it as A.
mucronata [fig.14]. But difficulties of identification were as nothing compared with trying to
photograph the tricky customer between the strands of a barbed-wire fence in the fading light of dusk.
To make matters worse, it was way down the bank beneath trees. Both of them took heaps of photos
and fortunately one, just one, proved of sufficient quality for publication here.
Of the two ferns we can say

little except to identify them.
They aren't our speciality, but
we know just enough about
pteridophytes to assign them to
genus level, then sort them out
to species level by looking up
those genera in our literature
and on the Internet. One is
Adiantum chilense [fig.15], the
other Cheilanthes hypoleuca
[fig.16]. Both were
photographed by Anita growing
together beneath cool rock
overhangs, and clearly did not
receive direct sunlight.
fig.15: Adiantum chilense, one

of the country's most plentiful
ferns. (30 Aug 2020. ARF)
fig.16: Another frequent pteridophyte in the Río

Aconcagua valley is Cheilanthes hypoleuca -
here a glaucous form.
(30 Aug 2020. ARF)

fig.17: Looking across the river at the private land to the E, where the new cistanthe was found.
(3 Nov 2020. JMW)
From this point, apart from a photo of the private sector, including its notice, taken across the stream
on the other side by John [fig.17], we continue perforce with descriptions and photos provided for us
by Carlos before he died. On crossing the river he continued to follow its upper course along the track
on the east bank. We know little directly about the flora of that sector, except that Tetraglochin alatum
[fig.18], an unusual shrublet of the Rosaceae, is fairly abundant there. He walked on towards the
mountains above [fig.19] through terrain covered by shrub cover and boulders, punctuated by
occasional Echinopsis chiloensis cacti looking like exclamation marks in the landscape. After
continuing for two and a half kilometres without seeing anything of interest, his attention was
suddenly caught by several dwarf plants with white flowers growing on an extensive bare, horizontal
strip of loose, sandy terrain. He'd discovered the type site of his Cistanthe celedoniana [fig.20]!
For the next two years, the last complete ones of his all too short life, Carlos continued on further up
the valley towards the snow-capped peaks above and found another population of 'his plant', adding
more photos of it [e.g. figs.67-70]. By great good fortune he also photographed a plant of the
accompanying flora in flower, Viola pusilla [fig.21], a small, yellow-flowered endemic annual rosulate
viola with loose foliage, which is plentiful throughout central and central northern Chile, with a
considerable extension to the south. It inhabits the littoral and inland areas to the Andean foothills at
1500 m. This detection by Carlos is particularly gratifying for the Watsons, whose principle focus of
study is the Andean violas, as it explains the presence in the valley of the monophagous fritillary
butterfly Yramea cytheris [fig.8]. Fritillary caterpillars only consume plants of that genus.
But Carlos left a further clue in a photograph he took there showing several other Andean herbs yet
to flower growing close to a small cistanthe [fig.22]. We've managed to identify one to species level,
the very variable Cristaria dissecta [fig.23]. It can be tall or short, glabrous, as here, or covered in
indumentum. Perennial calandrinias are familiar
enough to those who've visited South America or
read several of the many accounts of travels and
explorations there. A few are even in cultivation.
Much less well known are the several quite
different annuals of the genus, five of which
occur in Chile. We've been unable to fully identify
the one not yet flowering [fig. 22]. Cistanthe
grandiflora [fig.70] might be seen as a Gulliver of
the genus, whereas C. celedoniana is one of its
Lilliputians. We believe the small, glaucous-
succulent, innocent-looking, recently germinated
plant with broad leaves [fig. 22] cannot be
anything but the former. It's alarming to realise
that if this great dominant 'bully' were to colonise
the sandy stretch it would probably extinguish all
the small plants there, including C. celedoniana.
fig.18:Tetraglochin alatum is one of the shrubby

elements of the general cistanthe habitat.
(8 Aug 22013. Marijn van den Brink)
fig.19: Nearing the type location of Cistanthe celedoniana. (5 Oct 2014. Carlos Celedón)
fig.20: Cistanthe celedoniana at the type site on the day of its discovery. (30 Sep 2012. C. Celedón)
fig.21: The early flowering annual Viola pusilla, the only plant that Carlos saw immediately
accompanying Cistanthe celendoniana. (5 Oct 2014. Carlos Celedón)
fig.22: A small Cistanthe

celedoniana (in flower, far
right) and accompanying
taxa out of flower. (5 Oct
2014. Carlos Celedón)
fig.23: Cristaria dissecta, seen as a

glabrous form in leaf by the cistanthe, here
in its taller, hirsute form. (5 Sep 2020. JMW)
That covers all we know about the relevant

fieldwork of Carlos, but Christian can add
valuable detailed information and
illustrations of the flora that grows near, or
in the immediate vicinity of his location of C.
celedoniana in the Reserva Ecológica
[figs.24, 25], albeit very distant from Río
Colorado.
Slightly lower down from the cistanthe

populations, a most unusual member of its
genus was noted in flower. Malesherbia
fasciculata [fig.26] from an equally unlikely
herbaceous genus of the Passion flower
family ranges from Peru to Chile and
Argentina and is mainly Andean - with a few
along the central and northern temperate
Pacific coast. M. fasciculata looks even less
like a passion flower, or indeed like other
taxa of its genus, as its relevant unique
morphology consists of a tightly
compressed head of many very small
flowers.

fig.24: Cistanthe celedoniana habitat at the La Campana National Park.

(18 Nov 2020. Christian von Bohlen)
fig.25: A more
immediate view
of a La Campana
National Park
Cistanthe
celedoniana
habitat. (18 Nov
2020. C. von
Bohlen)
fig.26: Malesherbia fasciculata, a very

atypical member of its genus. They
usually have much larger, separate
flowers. (22 Aug 2018. JMW)
At the north-exposed lower site (965 m)

the Reserva shares with Río Colorado
the presence of an annual Calandrinia,
also not yet identified for certain, and five
more taxa besides. Montiopsis trifida
[fig.27] is the most plentiful and
widespread of its genus by far, but
despite inhabiting the length of Chile -
bar the extreme south - from near the
coast to 3300 m in the high Andes, it's
again endemic. In common with C.
celedoniana and most or all of that
species' immediate accompanying flora
it's an annual, as per the rather less
frequent congeneric species, M.
ramosissima [fig.28], another endemic,
which accompanies them both in the
Reserva locality.
The last three we know which grow in the immediately lower vicinity of C. celedoniana, again annual
and Chilean endemics, are far less widely known. All occur in the same three central regions, two of
them extending a bit beyond. Calycera sessiliflora [fig.29] belongs in the homonymous family, which
is restricted in distribution to the southern end of South America. Systematically, Calyceracaeae is
closely related to the composites and teasels, not least for its tightly bunched, multi-flowered
inflorescences. C. sessiliflora has a fairly limited elevational range from 700 to 1900 m.

fig.27: Montiopsis trifida, quite often seen by the authors in the general vicinity of their homes.
(22 Oct 2015. JMW)
fig.28: Montiopsis ramosissima, easy to tell from other species of its genus by the prostrate radiating
inflorescence 'spokes'. (Photo courtesy of Peter Peterson)
fig.29: Calycera sessiliflora, one of the less spectacular of the 'Andean cauliflowers'.
(Photo - Michail Belov)

fig.30:
Homalocarpus
dichotomus
(Apiaceae),
neatly midway
in stature
between
hemlock and
azorellas.
(Photo -
Michail Belov)
fig.31: Microphyes minima (Caryophyllaceae). This representative of a little-known genus looks

perennial. Sadly, it isn't. (Photo - Patricio Novoa)

A very dwarf and attractive species of the large family Apiaceae, Homalocarpus dichotomus [fig.30],
would surely appeal to alpine gardeners. Along with delightful little Microphyes minima [fig.31] of the
Caryophyllaceae, the final species known to the present authors which shares the immediate lower
habitat of C. celedoniana, it too reaches alpine levels. Microphyes makes one want to pick it up and
stroke it. Without doubt, if available it would be an even greater desideratum to add to the choicest
alpines in cultivation (sad it's only annual though). Curse all those who've put an end to responsible
(and we mean responsible) plant introduction! Would private vehicles be banned because there are
bad drivers and accidents, including when public figures are killed? Over time plants in general
cultivation have undeniably added greatly to botanical knowledge and significantly extended ex situ
conservation (Watson 2009).
fig:32: Argylia adscendens var. adscendens. A remarkable location of this usually Andean taxon at its
lower elevation limit. (17 Oct 2020. C. von Bohlen)
The higher site at 1603 m faces west. It possesses the flora of the lower site, but with the addition of
two perennials, both of which are more frequent and familiar at significantly higher elevations in the
main Andes to the east. Variable Argylia adscendens [fig.32] (Bignoniaceae) with its large broad
lobed, wide-tubed corolla, as well as the hummingbird adapted, tubular red amaryllid Zephyranthes
tenuiflora [fig.33] are further frustratingly mouth-watering desiderata.
fig.33: Handsome Zephyranthes tenuifolia is better known by its synonym Rhodophiala andina.
(17 Oct 2020. C. von Bohlen)
fig.33:
Type specimens
of Cistanthe
celedoniana
(A. Elvebakk)

Taxonomy (1)
Cistanthe celedoniana J.M. Watson, A.R. Flores & Elvebakk, sp. nov.
[figs.20, 33-61, 66]
Type: CHILE, Región de Valparaiso: Provincia de Los Andes, N side of Río Aconcagua Valley, 22 km
E of Los Andes, ca. 5 km NE of Río Colorado, 32º50'S 70º21'W, 1360 m, 30 Sept. 2012, leg. C.
Celedón, F& W 12801 (holotype CONC! isotypes CONC, SGO!).
Diagnosis:—The taxon described here differs from all other species of Cistanthe by its strongly and
regularly gibbose leaves, together with the combination of white petaloids, sepaloids equal in length
to the petaloids, and its regularly mottled seed indumentum.
Description:—Annual, rosette-forming subacaulous, glabrous herb, branched basally into 2–10 lax,
fertile shoots, forming plant 5–15 cm wide and up to 15 cm tall from fusiform tap-root to 10–12 cm
long and 3(–5) mm diam. Leaves thickly succulent, glaucous, margins sinuous; strongly gibbose,
gibbae 3–4 mm wide × 1–2 mm high formed pairwise, at times obliquely so, on adaxial surface only,
these developed on all leaves; base decurrent, subamplexicaul; apex rounded, tip acute to apiculate,
occasionally reddish brown in some allopatric populations. Rosette leaves 15–40 × 5–10 mm, simple,
linear-oblanceolate, acute; cauline leaves similar, but shorter and narrower. Stipules 3–4 × 2–3 mm,
amplexicaul, entire, triangular, apex acute, striated with black irregular lines, becoming hyaline.
inflorescence ca. 3-3.5 cm, thickened basally, dark maroon-red apically, at times branched, with 3–10
flowers in terminal, lax racemes; basal node sterile, 1-bracteose, 3-7 floral nodes with 2 unequal
bracts; peduncles arranged radially, 5–10 × 2–3 mm, at anthesis ascending to erect diurnally,
prostrate nocturnally. Pedicels to 6 mm, ascending to erect diurnally, prostrate nocturnally. Flowers
perfect. Sepaloids 5–6 × 6–7 mm, comprised of 2 unequal involucral bracts embracing the petaloids
in a form equivalent to sepals, non-keeled, strongly concave, broadly ovoid, apex acute with rounded
tip, intense green, strikingly and densely black-mottled and lined, partly along mid-vein, partly
irregularly; black markings very rarely absent, persistent. Petaloids 4–7 × 3–5 mm, 6–7, occasionally
as few as 5, arranged radially and symmetrically, free, imbricate basally, obovate, obtuse,
emarginate, with apical indentation 0.5 mm deep, pale greenish white with yellow base. Stamens
free, 7–14, 3 mm long; filaments yellow, gradually flattened towards 0.5 mm wide base; anthers 0.4-
0.5 mm, dorsifixed, yellow, pollen orange. Pistil superior, equal to or shorter than the stamens,
sincarpous. Ovary ca. 2mm long x1 mm diam., globose-ovoid, unilocular, placentation central,
greenish-yellow at anthesis; style 0.5-1 mm long, pale green; stigma opening into 3 oblong greenish
lobes, finely papillose. Fruit septicidal capsule 5-6 mm, with 19-23 seeds part-concealed by persistent
sepaloids, ovoid, dehiscing longitudinally as 3 papery valves, these pale brown, nerves well marked.
Seeds resembling a snail shell, 1–1.4 mm diam., lenticular to subglobose, weakly compressed,
outline subcircular to reniform, pale brownish yellow with outer circular marking radiating blackish
brown lines to margin, densely covered in minute transparent indumentum. Elaiosomes present, c.
0.3 mm wide, pale ochraceous.
Additional specimens studied, including of related taxa:—Cistanthe celedoniana J.M. Watson &
A.R. Flores. CHILE. Valparaiso Región, Provincia de Quillota, Palmas de Ocoa, Reserva Ecológica
Oasis de La Campana, sendero La Chusquilla, en terrano con suelo de maicillo, ladera exposición
norte, 32º56'28"S 71º01'52"W, 965 m, 5 Oct 2020, von Bohlen 2501 (paratypes CONC, SGO);
Cistanthe arenaria (Cham.) Carolin ex Hershk. CHILE, Coquimbo Region, Chungungo, 20 m, 15
Sept. 2008, M. Belov Z8226 (TROM); Cistanthe crassifolia (Phil.) Carolin ex Hershk. CHILE,
Atacama Region, Carrizal Bajo, 5–100 m, 30 Oct. 2008, M. Belov 1190 (TROM); Cistanthe.
grandiflora (Lindl.) Schlecht. CHILE, Maule Region, Laguna Maule, 1300–2100 m, 2 Sept. 2006, M.
Belov 0161 (TROM).

Distribution:— All known populations are confined to the Chilean region of Valparaiso. As seen by
the Watsons the type site and one other are situated in the Andean foothills beside the Río Colorado,
a tributary of the upper Río Aconcagua, on gently west sloping terrain at 1360 m and ca. 1500 m on
the north valleyside of the Río Colorado where it passes down through the Andean foothills. The
locations discovered by von Bohlen face north (lower) and west (upper), being ca. 70 km distant to
the SSW of Río Colorado and situated in the in the hills of the Reserva Ecológica de La Campana at
965 and 1603 m on a gradient of 30º and with north and west exposures respectively.[figs.2-4]
General and immediate flora at cistanthe habitats:— The overall vegetation cover of the two
known locations is classified by Gajardo (1994) as Matorral Esclerófilo (Mediterranean esclerophyll
shrub land), and consists of biodiverse and widely dispersed low xerophytic taxa, including Baccharis
paniculata (Asteraceae), reduced forms of Lithrea caustica (Anacardiaceae), Kageneckia oblonga
(Rosaceae), Podanthus mitique (Asteraceae), Tetraglochin alata (Rosaceae) [fig.18], Trevoa trinervis
(Rhamnaceae) and others. Taxa of Haplopappus and Senecio (both Asteraceae) have not been
identified to species level. The cactus Echinopsis chiloensis was also observed.
fig.35: A pressed flower of the type specimen of Cistanthe celedoniana. (A. Elvebakk)
Both populations at Río Colorado and the two at the Reserva inhabit extensive almost bare stretches
of deep, fine to coarse grained and compacted but readily permeable sand. At times it is interspersed
with occasional small rocks. The cistanthe and its accompanying flora were scattered over this
terrain. The latter consists of Calandrinia sp. (Montiaceae) [fig.22], possibly another Cistanthe sp.
[fig.22], Cristaria dissecta (Malvaceae) [figs, 22, 23], and Viola pusilla (Violaceae) [fig.21] at least,
according to what can be seen on the photographs of Celedón. The accompanying Reserva flora has
been surveyed and identified more thoroughly, and as seen consists of Calandrinia sp (Montiaceae),
Calycera sessiliflora (Calyceraceae) [fig.29], Homalocarpus dichotomus (Apiaceae) [fig.30],
Malesherbia fasciculata (Passifloraceae) [fig.26], Microphyes minima (Caryophyllaceae) [fig.31],
Montiopsis ramosissima [fig.28] and M. trifida (Montiaceae) [fig.27] at the lower site, with the addition
of Argylia adscendens [fig.32] and Zephyranthes tenuiflora [fig.33] at the second higher up. There is a
significant difference in the composition of the relevant floras at both the above locations, and this
may well apply to any others found in future, which are equally distant.
Phenology:— Based on observations in the wild and in cultivation, we deduce that anthesis
commences in September and continues into November, with seed dispersal following at a short
interval after each individual flowering.
Etymology:— This species is named in memoriam for our late friend and amateur naturalist Carlos
Celedón (1969–2015), and also commemorates his significant botanical achievements. He was an
indefatigable field explorer who discovered the present new species as well as another and provided
us with plant material, information and photographs. Regrettably he passed away at much too early
an age.
Considered conservation status:— The two populations known to the authors contain relatively few
individual plants. The type location apparently comprised ca. 50 when visited, and the nearby higher
second population fewer, perhaps no more than 30, both populations close-set (C. Celedón, pers.
comm.). Its known extent of occurrence (EOO) would be about 2-3 km, its total area of occupancy
(AOO) being linear, can scarcely be any wider. The number of observed plants was therefore
somewhat fewer than 100 individuals.
The Reserva Ecológica de La Campana location, 70 km distant, consists of two closely separated
populations, the lower of which is estimated to contain at least 40 to 50, and the higher 25 to 30
individuals of C. celedoniana, (C. von Bohlen pers. obs.). Although situated in a private nature
reserve, there is no guarantee of assured, permanent protection.
Given the close similarity of these statistics at both these very widely separated main locations,
perhaps adding up to slightly more than 200 individuals between them over an EEO of 70 km, it
seems reasonable to assume that any undiscovered intermediates or extensions of the species'
range would be similarly composed and disposed. Accordingly, by applying IUCN Red List criteria
and guidelines (IUCN 2012), a conservation status of CR (critically endangered) based on present
knowledge is proposed here.
Seed morphology
All Cistanthe species studied have a black, glossy surface with irregularly hexagonal and weakly
convex areoles, separated by a tissue forming a reticulate network of distinct and strongly convex
lines ca. 4 µm wide. In C. celedoniana [figs.52-61, 66] the 15–20 µm wide areoles consist of a central
plate of concentric structures, surrounded by transparent tissue, resembling a fried egg. The central
part of a scale produces a trichome, first papillose and brown, later conical-spathulate, constricted at
the base and flattened, then becoming white-transparent when fully developed in the elongated part.
These trichomes form dark patches of short papillose trichomes, alternating with other patches where
the trichomes have white-transparent elongations. In C. grandiflora [figs.62, 63, 66] the areoles are
20–40 µm wide with distinctly radiating patterns. The trichomes are digitiform, whitish and
transparent, pale brown only at the base where some have weak dilations. Trichomes of C.
crassifolia [figs.64-66] are similar to those of C. grandiflora. Those of C. arenaria are very different
insofar as they possess coralloid branching.
Pollinator activity, pollination and pollen dispersal
In the field at Río Colorado ants were observed to feed frequently on nectar in the flowers. Some
arrived at plants with nothing adhering to them [fig.67], but as they moved around the flowers feeding
[fig.68] they became progressively covered in pollen grains, which, as they exited, covered their legs,
head, thorax and abdomen [fig.69]. On entering flowers, some already had a covering of pollen,
which clearly derived from visiting another corolla (C. Celedón pers. comm.). Thus cross-pollination
of the cistanthe is achieved.
Ant pollination was reviewed by Del-Claro and coauthors (2019), who indicated that it is a rare but
partly overlooked exclusive insect and plant interaction, involving plants with near-ground
inflorescences, often with white flowers, and concentrated on relatively arid habitats.
Although this species is pollinated by ants, as thoroughly observed [figs.67-69], it is not exclusively
so. Two beetles were also photographed visiting the flowers [fig.70] at the Río Colorado upper
location. By contrast, no ant activity at all has been observed at the Reserva Ecológica de La
Campana, whereas other unidentified insects were noticed there on the upwards-facing flowers of
erect stems (C. von Bohlen obs.). These flowers are evidently positioned to attract passing
arthropods on the wing.
Diurnal plant tropism
Cultivated specimens transplanted from in situ seedlings flourished and flowered freely,
synchronously with their cycle in the wild, but presented some unexpected diurnal rhythms as
observed several times. From late afternoon onwards until noon the following day stems were
declined [figs.36, 39, 40, 43]. However, by 2 pm they had ascended [figs.42, 44]. A similar pattern
can be noted in photographs taken in the wild [figs.20, 39, 40, 41]. Available seeds from the ex-situ
pot plants did not germinate, so the species is no longer in cultivation to provide further observations.
Plant stems as shown in [figs.36-40, 43] may perhaps partly at least involve - or have evolved from -
myrmecotrophism, defined as plant movement to attract ants. The horizontal and early morning
position of the branches at ground level would place the conspicuous mottled contrast pattern of the
sepaloids close to the ground and clearly visible to ants, attracting them initially to the easily
accessible corollas as these are opening under the influence of light (photonasty). An erect or sub-
erect branch position with its conspicuous flowers later in the day would then cater to a normal
selection of pollinating insects, although ants do in fact continue to visit then (C. Celedón, obs).
These foliar and floral actions have been recorded from the wild as well as cultivation (Watson &
Flores obs.), so cannot be attributed to some artificial effect of the latter. The lowering of the shoots
and closing of the flowers is a nocturnal (nyctinastic) circadian rhythm induced by the onset of dusk
and possibly also by reduction of air temperature.
Minorsky (2018) reviewed existing hypotheses explaining the evolutionary function of this tropism and
added the idea that it might be an adaptation to protect plants from nocturnal herbivores. That could
offer three possible benefits: making the plant as inconspicuous as possible; removing its stems from
the erect position favoured by mammalian herbivores; and increasing the possibility of revealing
arthropod herbivores to their predators. The phenomenon may well serve both these purposes at
least. Interestingly, the Watsons deduced the same possibility from pure observation. Well organised
future investigation is needed to discover what lies behind this remarkable adaptation.

Left, fig.36: Wild-transferred plants in the morning prostrate position with closed flowers. (13 Nov
2014. JMW) Right, fig.37: A wild C. celedoniana, totally prostrate with closed flowers in the evening
of a dull day. (4 Oct 2020. C. von Bohlen)
Left, fig.38: Cistanthe celedoniana in habitat at mid-day, part prostrate still but with flowers open.
(4 Oct 2013. Carlos Celedón) Right, fig.39: Successful cultivation of plants from the wild, here rising
from the prostrate posture. (12 Oct 2014. JMW)
Far left, fig.40: Wild
collected individuals
with flowers open in
the plant's prostrate
phase.
(29 Oct 2014. ARF)
Left, fig.41: A wild
plant in the erect
daytime position.
(17 Oct 2020.
C. von Bohlen)

fig.42: Cultivated individual transferred from

the wild in fully erect posture.
(29 Oct 2014. ARF)
fig.43: Cistanthe celedoniana. A collected plant in

cultivation showing depressed overnight pre-flowering
position at 11.48 a.m. (13 Nov 2014. JMW)
Left, fig.44: Cistanthe celedoniana. The same cultivated individual (as in fig.43) in erect flowering
position at 2.36 p.m. on the same day. (13 Nov 2014. JMW)
Right, fig.45: Cistanthe celedoniana in habitat. Foliage presentation and gibbose structure.
(30 Sep 2012. Carlos Celedón)
fig.46: Cistanthe celedoniana. Lamina, showing gibbosities and their tendency to alternate at times.
(7 Nov 2014. JMW
fig.47: Cistanthe celedoniana in habitat, the sepaloids typically striated. A less common cruciform
corolla is evident. (30 Sep 2012. Carlos Celedón)

fig.48: Inflorescence of C. celedoniana from La Campana National Park the Reserva Ecológica
de La Campana. Note smaller flower. (15 Oct 2020. C. von Bohlen)
fig.49: Cistanthe celedoniana in habitat. Fully open and unfolding corollas, the latter showing the
similar lengths of sepaloids and petaloids. (30 Sep 2012. Carlos Celedón)

fig.50: Cistanthe celedoniana in habitat. An atypical rare specimen with unmarked sepaloids.
(4 Oct 2013. Carlos Celedón)

fig.51: A dense inflorescence of C. celedoniana showing sepaloid marking and an open capsule with
seeds. (15 Oct 2020. C. von Bohlen)
fig.52: Cistanthe
celodoniana seeds.
(Photo - Mari
Karlstad)

Seed vectoring
The concurrence of elaiosomes and mottled sepaloids in most species of Cistanthe, as compared
with absence of that morphology in neighbouring Montiaceae genera, is more probably a
myrmecochory syndrome. Whether or not they have pollinated Cistanthe taxa in flower, it is likely that
ants represent the main agents for dispersing seeds of them all. This may also be the case with the
hirsute seeds of most species of the section. As outlined by Gómez et al. (2005), surface seed
structures functioning as a handle may aid dispersal after the elaiosomes have been consumed by
the ants, the latter a process which might also promote scarification, thereby improving germination
(Hughes & Westoby 1992).
Cistanthe celedoniana apparently also has a further adaptation to myrmecochory as represented by
its distinctly mottled seeds [fig.52]. This pattern is obtained by short brownish trichomes being
extended into white longer forms distributed in regularly alternating patches [figs.57-61]. Cistanthe
celedoniana would evidently be an excellent subject for future in-depth studies on interactions
between plants and ants.
[The following nine images (figs.53 – 61) are of seed of Cistanthe celedoniana, µm (micrometre) by
Arve Elvebakk.]
fig.53: (a) Cistanthe celedoniana
fig.54: (b) Cistanthe celedoniana
fig.55: (c) Cistanthe celedoniana
fig.56: (d) Cistanthe celedoniana

fig.57: (e) Cistanthe celedoniana
fig.58: (f) Cistanthe celedoniana
fig.59: (g) Cistanthe celedoniana
fig.60: (h) Cistanthe celedoniana
fig.61: (i) Cistanthe celedoniana

Left, fig.62: (j)Cistanthe grandiflora, µm (micrometer) photograph of seed. (A. Elvebakk)

Right, fig.63: (k) Cistanthe grandiflora, µm (micrometer) photograph of seed. (A. Elvebakk)
Left, fig.64: (l) Cistanthe crassifolia, µm (micrometre) photograph of seed. (A. Elvebakk)
Right, fig.65: (m) Cistanthe crassifolia, µm (micrometre) photograph of seed. (A. Elvebakk)

fig.66: The same figures with bars showing the scale in µm (micrometres) ⁓ a, d, f, h = 200 µm ⁓ b,
c, g, i‒m = 30 µm. (A. Elvebakk)

Taxonomic context with selected references
Shortly after its inconspicuous start as just two species, the genus Cistanthe Spach (1836) was soon
sunk into synonymy with Calandrinia Kunth (1823) and treated as such by all later important studies
for almost 150 years. These include, inter alia, Barnéoud in 1846 (by implication of taxa cited, as he
did not in fact mention the name Cistanthe); Reiche (1897) (still the best monograph of the Chilean
species); Pax and Hoffmann (1934); Añon Suarez in her 1953 monograph of Calandrinia in
Argentina; vol. 2 of Flora de la Cuenca de Santiago de Chile by Navas (1976); Marticorena &
Quezada (1985); Peralta (1988); and Marticorena (1992).
A revision process was initiated by Carolin (1987), who highlighted the high heterogeneity of
Calandrinia. Based on the evaluation of many morphological and anatomical characters and a
cladistic analysis, he proposed splitting Calandrinia into five genera. He himself continued to treat
Calandrina collectively though, but with indications of proposed future generic affiliations of the major
groups within existing Calandrinia sens. lat.
Subsequently, Cistanthe was resurrected as a separate genus by Hershkovitz (1990). He supported

the conclusions reached by Carolin (1987), but also included the North American genus Calyptridium
with its eight species as a separate section of Cistanthe, and the monotypic section Strophiolum,
previously assigned to Lewisia, as another section. This treatment is still followed by Kelly, Miller and
Packer in the Flora of North America in its present 2019 version. Hershkovitz (1990) also included
two small South American alliances, the genus Philippiamra Kuntze and Calandrinia sect.
Amarantoideae Reiche as new sections within Cistanthe. He dealt with all five sections more
thoroughly in 1991. The largest group within the genus was Cistanthe sect. Cistanthe, which, after
recombinations made by Hershkovitz in 1991 and subsequently by others up to 2008, amounted to
27 taxa. Peralta & Ford-Werntz (2008) and Peralta (1993) augmented the total with another five
species in the remaining sections in South America.
Former support for the monophyly of Cistanthe sens. lat., including its sections, was found to be
weak by Hershkovitz and Zimmer in 2000, and already in 1999 the monotypic section Strophiolum (B.
Mathew) Hershk. was treated as the new genus Lewisiopsis by Govaerts. In 2015 Calyptridium
Nuttall was resurrected as a separate genus, rendering remaining Cistanthe taxa less
heterogeneous, except that they rejected Lewisiopsis as a separate genus.
Hershkovitz recently produced a monograph of the Montiaceae (Hershkovitz 2019). He now defined
Cistanthe as only including its previous sect. Cistanthe, which he divided into a more narrowly
defined sect. Cistanthe, consisting primarily of perennial species, and the new Cistanthe sect.
Rosulatae (Reiche) Hershk., mainly of annuals. The latter was divided into two subsections, subsect.
Rosulatae (Reiche) Hershk. and subsect. Thyrsoideae Hershk. He considered the genus to total at
least 38 species, including six new recombinations and his own Cistanthe subspeciosa Hershk. He
has also recently described Cistanthe philhershkovitziana from the northern Mediterranean zone of
Chile, and named it in honour of his father. Another to be recently described, Cistanthe floresiorum
J.M. Watson [fig.72], from a pass in the Atacama Region of Chile to the north of La Serena, is named
after the family of the author‘s wife, principally her parents, who discovered it at the type locality
(Watson 2019).
The information presented by Peralta & Ford-Werntz (2008) shows that 11 species are endemic to
the northern Atacama Desert of Chile. Of these, Cistanthe longiscapa (Barneoud) Carolin ex Hershk.
is a primary contributor to the extensive magenta coloration of the Flowering Atacama Desert
phenomenon. Another four species also occur both there and in adjacent provinces of Argentina,
while five species are endemic to the mediterranean zone of Central Chile. Finally, four much smaller
and more depressed species are Andine, occurring between altitudes of 2000 and 5000 m, and are
also again shared by Chile and Argentina.

Cistanthe, with such a strong evolutionary centre in northern Chile, is still badly in need of renewed
in-depth studies, and to that end Hershkovitz (2019) cited its section Rosulatae as ‗woefully
inadequately‘ known. Without doubt, for most interested parties that assessment could equally be
applied to sect. Cistanthe as a whole! As an example of this unfamiliarity, the 2019 updated
presentation of Cistanthe species endemic to Chile by the Royal Botanic Garden at Edinburgh
included 24 species when consulted, but only six were illustrated.
The present work describes another newly discovered, distinct Cistanthe species, at present known
from two general localities in Valparaiso Region, central Chile. We also include a recombination of
another species, originally intended for a now cancelled guide to the Atacama Flowering Desert,
which - as Vol. 2 - was to have covered the dicotyledons of the country's northern Pacific coast and
adjacent interior lowland, the monocotyledons having been already published (Hoffmann et al. 2015).
Systematic relationships
Cistanthe is readily separated from its neighbouring genera Calandrinia and Montiopsis, all now
included in the family Montiaceae (Nyffeler & Eggli 2010); e.g. by a combination of its strongly
succulent leaves, its characteristic black blotched and striated sepaloids, and its seeds, which are
mostly tomentose and strophiolate with elaiosomes, and not compressed laterally. Cistanthe
celedoniana differs from all other species by its strongly gibbose leaves, which would probably make
the species attractive in any specialist succulent collection.
The only Cistanthe species which at times bears a certain resemblance to C. celedoniana, is C.
arenaria [fig.75] as presently interpreted; i.e. extremely polymorphic (Ford-Werntz & Peralta 2002,
inter alia). This widespread species from the same biogeographic region is usually strongly erect, and
with different foliage, but some of its forms have slightly but significantly gibbose adaxial leaf
surfaces, as can be seen in fig.75. Its flower colour is primarily magenta, although white-flowered
forms do occur occasionally in most magenta species. However, the long sepaloids of C.
celedoniana, equally long as the petaloids, present a key character separating it from all forms of C.
arenaria. It should be borne in mind that Hershkovitz (2019) advocated a revised interpretation of C.
arenaria a topic he then planned to deal with in more detail later.
The annual life cycle of C. celedoniana indicates that it belongs in section Rosulatae. Hershkovitz
(2019) did not cite any other characters than this when defining the section, which comprises several
of the former sections of Calandrinia sensu Reiche (1998). Hershkovitz (2006) indicated that the
other recently described new annual species C. floresiorum (Watson 2019) resembled a reduced
version of perennial C. grandiflora. However, C. floresiorum had already been analysed genetically
by him (Hershkovitz 2006) as ‗Cistanthe sp. 02 80‘ and confirmed then to be a species in sect.
Rosulatae (Reiche) Hershk. as was proposed by Watson (2019), also as based primarily on its
annual life cycle.
Those of Nyanano in 1988 (but with no Cistanthe species included) and Peralta (1996) are the only
previous SEM based studies of seed morphology in the families Montiaceae and Portulacaceae,
apart from illustrations of seeds of the four Patagonian group Calandrinia species. These were
presented by Elvebakk et al. (2015) in their paper which included the new species C. ranunculina.
Cistanthe grandiflora and C. crassifolia are related species, although the latter was not treated as a
separate taxon of Cistanthe until very recently (Hershkovitz 2019). Both being sect. Grandiflorae
members, they share an analogous seed micro-morphology [figs.62-66], i.e. dominated by long,
transparent and finger-like trichomes, sometimes with weak basal swellings. Cistanthe picta (Arn.)
Carolin ex Hershk. [fig.73], Cistanthe frigida, (Barnéoud) Peralta and Cistanthe humilis (Phil.) Peralta
[fig.74], deviating and perennial Andine members of sect. Rosulatae (Hershk.), were shown to have
short outgrowths on the seed exterior, in the first club-shaped (‗en forma de porra‘), in the second like
a glove finger (‗en forma de dedo de guante‘). C. humilis has not been analysed. In C. arenaria
[fig.75] they are coralloid and very different. Concerning the flattened trichomes with basal
constrictions possessed by C. celedoniana [figs.57, 58], they most resemble those of sect. Cistanthe.
The radiating structure of the seed scale plate of C. grandiflora [fig.64] is surprisingly different from
the ‗fried egg‘ structure with concentric structures in C. celedoniana [figs.57, 58]. So far, these
microscopic characters have been studied in too few species to reveal patterns within Cistanthe.
However, the high diversity in seed ultrastructure shown so far appears to be a promising analytical
character in Montiaceae/Portulacaceae taxonomy. The morphological differences between the
studied representatives of sect. Grandiflora vs. sect. Cistanthe should be investigated further to settle
whether they might represent new synapomorphies for these two sections.
fig.67: Ant entering Cistanthe celedoniana flower at 11.21.57 a.m. (5 Oct 2014. C. Celedón)
fig.68: Ant now covered in pollen while moving and feeding. 11.24.32 am (2 mins 35 seconds later).
(5 Oct 2014. C. Celedón)
fig.69: Ant dusted with pollen leaving flower at 11. 25.53. Duration of stay was 3 minutes 36 seconds.
(5 Oct 2014. C. Celedón)
fig.70: Beetles on Cistanthe celedoniana flowers. (30 Sep 2013. C. Celedón)

fig.71: In contrast to Cistanthe celedonium, C. grandiflora is a vigorous, large-flowered perennial.

(9 Oct 2010. JMW)

fig.72: F.& W. 11750 Cistanthe floresiorum, illustrating the cymose presentation. Pajonales Pass.
(19 Oct 2008. JMW)
fig.73: Cistanthe picta is low growing, like C. celedoniana, and sometimes white-flowered, but
perennial. (29 Jan 2009. Michail Belov)

fig.74: The tiny, always white-flowered Andean, Cistanthe humilis, is also perennial.
(10 Feb 2011. JMW)
fig.75: M.R. 8838 Cistanthe arenaria. Note gibbose leaves of this form as in C. celedoniana.
(14 Oct 2014. Marcelo Rosas)
Concluding thoughts
It may seem quite amazing that such a distinct local endemic species has remained undescribed until
now, growing as it does a mere 60 km away from Santiago at the type location, and in a widely
populated geopolitical region which had apparently been well-explored botanically. However, there
are several other examples of recent discoveries of striking plants from this area of central Chile with
its very high level of endemism, e.g. the beautiful Alstroemeria piperata A.R. Flores & J.M. Watson
(Watson et al. 2018) [fig.79] and the unique and magnificent Viola regina (Watson & Flores 2020)
[figs.80, 81], both also involving Carlos. No less relevant in this context are many rediscoveries,
including of famous Tecophilaea cyanocrocus in the Santiago Andes (2001).
Carlos Celedón was among those amateur wildflower

enthusiasts who've played an important role in
making known such rare plants, often threatened by
land-use and other factors. Remember too: he did it
all for 'love', and didn't receive a peso, penny or dime
for his efforts. As a lay naturalist he did not have or
need a C.V. to boost a career either.
It is to be hoped this publication may initiate further

searches for Cistanthe celedoniana, perhaps
interconnecting the present widely separated
populations as known and even finding other
undescribed plants. We feel it's appropriate to end the
review of his cistanthe with portraits of Carlos [figs.76,
83], and the five species [figs.77-82] that have been
contributed to the world of science thanks to his
devotion and tireless energy.
fig.76: Carlos in head-down plant hunting posture.

(18 Aug 2013. JMW)
Taxonomy (2)
A new combination
Cistanthe sitiens (I.M. Johnst.) A.R. Flores and J.M. Watson, comb. nov.
Basionym:— Calandrinia sitiens I.M. Johnst., Contrib. Gray Herb. 85: 35. 1929.
This rare species is only known from the Tocopilla area (22º05'S 70º12'W) in the Atacama Flowering
Desert of Antofagasta Region in northern Chile. It was reported and illustrated by Finger & Tellier
(2010), as Calandrinia sitiens, but is clearly a Cistanthe species. This was evident to the authors of
the new combination who intended to publish it in the aforementioned and cancelled field guide.
Instead, that taxonomic change is effected here.

―The five species that have been contributed to the world of science thanks to his devotion and
tireless energy.‖
fig.77: The yet unpublished Oxalis ranchillos, found by the Watsons at Carlos's original site.
(24 Oct 2015. ARF)
fig.78: Originally Carlos took us to the 'long lost' Viola chamaedrys of Leybold. So began the
flowering of our co-operation. (8 Sep 22013. JMW)
fig.79: Alstroemeria piperata, a little beauty also found by John and Anita at the Viola chamaedrys
site. (20 Dec 2015. ARF)
fig.80: Carlos's most exciting discovery, Viola regina, 'Queen of the Violas'.
(28 Nov 2013. C. Celedón)
fig.81: Edited photograph of the unique and unmistakable Viola regina flower. (C. Celedon)
fig.82: Cistanthe celedoniana (17 Oct 2020. C. von Bohlen)

Acknowledgements
We owe the late Carlos Celedón immeasurable posthumous gratitude for sharing all his information
and photographs of the new species with us. Local landowners of the Río Colorado area also
graciously allowed him to pass across their private land to reach its habitat. John and Anita
particularly wish to thank two staff members of the company which owns the land across the Río
Colorado, who casually met them and then drove them in their pick-up along the track taken by
Carlos. They have expressed the wish to remain anonymous. Marcelo Rosas of Coquimbo kindly
supplied a photograph of Cistanthe arenaria. Mari Karlstad, UiT – Arctic University of Norway, was
responsible for the seed illustration shown as [fig.52]. Others whose photos illustrate this presentation
are acknowledged in the captions. Tom-Ivar Eilertsen, gave us permission to use the Advanced
Microscopy Core Facility (AMCF) at the University of Tromsø, the Arctic University of Norway, where
the SEM equipment was operated by Augusta Hlin Aspar Sundbø.
fig.83: A parting shot of Carlos when he was prepared for a short exploratory excursion at Los
Ranchillos. (30 Nov 2014. JMW)

References
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Del-Claro, K., Rodriguez-Morales, D., Calixta, E.S., Martins, A.S. & Torezan-Silingardi, H.M. 2019.
Ant pollination of Paepalanthus lundii (Eriocaulaceae) in Brazilian savanna. Ann. Bot. 123: 1159–
1165. doi: 10.1093/aob/mcz021.
Elvebakk, A., Flores, A.R. & Watson, J.M. 2015. Revisions in the South American Calandrinia
caespitosa complex (Montiaceae). Phytotaxa 203: 91–23.
Finger, K. & Teillier, S. 2010. Contribución al conocimiento de la flora endémica de Taltal y Paposo,
Región de Antofagasta (II), Chile. Chloris Chil. 13, 2. http://www.chlorischile.cl.
Ford-Werntz, D. & Peralta, I. E. 2002. Cistanthe. In: Eggli, U. (ed.), Illustrated handbook of succulent
plants: dicotyledons, vol. 2: 380–386. Springer Verlag, Berlin, Heidelberg & New York.
Gajardo, R. 1994. La vegetación natural de Chile. Editorial Universitaria, Santiago, 165 pp.
Gómez, C. Espadaler, W & Bas, J.M: 2005. Ant behaviour and seed morphology: a missing link of
myrmecochory. Oecologia 146: 244‒246. doi: 10.1007/s00442-005-0200-7.
Hershkovitz, M.A. 1990. Nomenclatural changes in Portulacaceae. Phytologia 68: 267- 270.
Hershkovitz, M.A. 2019. Systematics, evolution, and phylogeography of Montiaceae
(Portulacineae). Phytoneuron 2019-27: 1–77.
Hoffmann, A.E., Watson, J.M. & Flores, A.R. 2015. Flora silvestre de Chile: cuando el desierto
florece volumen 1: monocotydedones y otros taxones. Fundación Claudio Gay, Santiago de Chile.
261 pp.
Hughes, L. & Westoby, M. 1992. Fate of seeds adapted for dispersal by ants in Australian sclerophyll
vegetation. Ecology 73: 1285–1299.
IUCN. 2012. International Union for Conservation of Nature red list categories and criteria: Version
3.1. Second edition.
Minorsky, P.V. 2018. The functions of foliar nyctinasty: a review and hypothesis. Biol. Rev.
Cambridge Phil. Soc. 94: 216-229. doi:10.1111/brv.12444.
Nyffeler, R. & Eggli, U. 2010. Disintegrating Portulacaceae: a new familial classification of the
suborder Portulacineae (Caryophyllales) based on molecular and morphological data. Taxon 59:
227–240.
Peralta, I.E. 1993. Distribución del género Cistanthe Spach (Portulacaceae) en Sudamérica.
Parodiana 8: 153–158.
Peralta, I.E. & Ford-Werntz, D.I. 2008. Portulacaceae. In: Zuloaga, F.O., Morrone, O. & Belgrano,
M.J. (eds.), Catálogo de las plantas vasculares del Cono Sur (Argentina, Sur de Brasil, Chile,
Paraguay y Uruguay). Dicotyledoneae: Fabaceae (Senna-Zygia) Zygophyllaceae 3. Monogr. Syst.
Bot. Miss. Bot. Gard. 107: 2796–2817.
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Watson, J.M. 2019. Flores means flowers ... of course! A new Cistanthe species (Montiaceae) from
Chile's springtime north. IRG 109: 2–42.
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Scot. Rock Gard. Club 133: 97-101
--- World of Iris ---
The King of Reticulata By Panayoti Kelaidis
Sometimes we get lucky. In March of 1987 I flew to Toronto for the first time for a horticultural
presentation at Edwards Gardens (now the Toronto Botanical Garden). Although my specialty has
always been alpine plants and rock gardening, I have a strong interest in the genus Iris, and I have
been a long time member of SIGNA (Species iris Group of North America) - one of the branches of
the American Iris Society. In their newsletters I had noted that Alan McMurtrie had been growing and
writing about species iris. I contacted him before my trip thinking perhaps he might have a few Iris to
see even early in the season during my visit. All this was conducted by mail and post offices back
then: remember, this is before email and the internet invaded our lives!
Alan McMurtrie in 1987: photo P. Kelaidis
I rented a car while in Toronto and devoted

a day to driving out to the suburb where
Alan lived and fortunately it was a sunny
day and sure enough, he had lots of irises
blooming—many of them for the first time:
he‘d taken two collecting trips to Turkey the
years prior to my visit, and I saw for the first
time wild collected specimens of Iris
danfordiae and Iris histrioides blooming—
altogether different from the plants we grow
under these names in our gardens. I had
come to see species, but Alan‘s talk was all
about his plans one day to hybridize plants
in the Iriodictyon family of the genus
(roughly a dozen species of plants we
loosely refer to as ―the reticulata section‖). I
listened to his rather grandiose descriptions
of what he had in mind at the time and I only
wish I‘d had recorded what he said on a
tape recorder. Although I thought what he
was describing was rather far-fetched, I
listened patiently. Little did I suspect the
reality of what Alan went on to achieve in
the years after my visit have far exceeded
anything he might have imagined in 1987.
It is all lovingly documented in amazing
detail on his enormous and wonderful
website: reticulatas.com.
Alan was employed at the time I visited him (and for many years thereafter) as an electrical engineer
working for the Canadian Power company - a job with great responsibility requiring specialized skills.
He was already married and had two sons who also occupied a great deal of his time and energy. I
find it remarkable that in his spare time he was able to create from scratch the most ambitious
hybridization program that I am aware of in the entire realm of hardy bulbs, crossing hundreds and
ultimately producing thousands of seedlings, the best of which he has had the vision and business
skills to market in the Netherlands where he has partnered with many Dutch growers to ultimately
produce many of his best hybrids for the retail market.

Iris sophenensis x Iris danfordiae, photo Alan McMurtrie
The year of my visit he began his first wide crosses between the fertile wild Iris danfordiae and Iris
sophenensis that provided his first major breakthrough in his breeding. After that beginning, his
program has literally sky-rocketed with breakthroughs and new colors.
Alan and I stayed in contact in subsequent years, and I followed his career with great interest. I was
dazzled when he published pictures of his first wide crosses where the dramatic orange, bronze and
multicolored iris flowers were like nothing I‘d ever seen before. I simply had to get my hands on some
of these bulbs.
Alan realized that the semi-arid steppe climate of Denver might especially suit his hybrids. Over the
last decade Denver Botanic Gardens has purchased large quantities of McMurtrie irises directly from
Holland to grow in our collections and also to sell at our large autumn plant and bulb sale (which this
year sold nearly $100,000 in bulbs and plants). The McMurtrie iris offerings have become a major
magnet for plant connoisseurs at this sale, and hundreds of Denver area gardeners are now enjoying
these in their private gardens.
I have been lucky to be able to obtain a large number of Alan‘s most recently marketed hybrids which
I‘ve grown in various parts of my home garden. The pictures accompanying this piece show how
beautiful they can be in a rock garden setting. In Denver they grow well in any well drained soil -

although I find they seem to like soils that have some loam rather than a sterile scree. They do need
moisture in the springtime (when Denver often has frequent, heavy snowfall in late winter), I have had
some of Alan‘s hybrids be snowed upon several times in a single spring: they are so sturdy they can
go through two or three snows and keep on blooming!
Iris x denticulata ‗Happiness‘ in the snow.

Each of Alan‘s hybrids has its own personality - but I have to admit that when ‗Mars Landing‘ first
bloomed for me, I had a sort of shiver of amazement: it was even stranger and more wonderful than
I‘d imagined! But ‗North Star‘ or ‗Sea Breeze‘ have every bit as much panache. I want them all!
In our sunny, steppe climate we can have bulbs blooming throughout the winter months: snowdrops
for instance bloom from October and November (Galanthus peshmenii and G. elwesii var.
monostictus) and by December and January the ―spring‖ crocus are already emerging and starting to
bloom. Crocus, Cyclamen, Sternbergia and Colchicum all bridge the fall to spring season—effectively
shortening our otherwise long winter season. Although we do not yet have autumn blooming Iris, the
Iriodictyon section are among the earliest bulbs to open in the new year -occasionally during a warm
spell in January we have had Iris danfordiae open a flower, but by mid February there are many
reticulatas starting to bloom in the warmest microclimates. Late February to mid March is usually the
peak season (invariably interrupted by two or three snowstorms - although the snow usually melts
quickly that time of year). There are usually a few reticulata section iris still blooming into the first
week of April - giving them almost three months when there is little competition in the rock garden
except for a few other ―minor‖ bulbs.
Left:
Iris ‗Mars
Landing‘
Right:
Iris ‗Sea
Breeze‘
Iris ‘Painted Lady‘
Each year there seem to be a few more McMurtrie irises

showing up in specialist bulb catalogues, and his older
hybrids are starting to become ―bread and butter‖ - which
is to say standard plants even in the mass market bulb
catalogues.
Ed.: Alan McMurtrie has been awarded the British Iris

Society's Foster Memorial Plaque (2010) and their
Hybridisers Award (2016) as well as the
American Iris Society's Hybridiser Award (2019).

Iris ‗Happiness‘ – this time

in full flower in sunshine.
They are comprising a

more and more important
element in my personal
garden - they are the stars
of the winter garden for me
along with Galanthus,
Adonis and Helleborus. But
the McMurtrie iris have far
more flamboyant colors
and a huge range of
variability. If you can
succeed with the old Dutch
selections like Iris reticulata
‗Harmony‘ or ‗J.S. Dijt‘ I
know you will welcome
Alan McMurtrie‘s
remarkable new
assemblage of rock garden
gems.
Iris 'Sunshine' Iris reticulata 'White Caucasus' and Tulipa humilis

Iris 'Finola'
Iris ‗Katharine‘s
Gold‘

Iris ‗North Star‘

Iris x reticulata 'Eyecatcher' and ‗Blue Hill‘
Above right: Alan

speaking at the
SRGC Early Bulb
Event in 2016.
Left:
A selection of his iris
hybrids at an RHS
London Show where
Alan McMurtrie was
able to chat with
members of the
public.

Cistanthe Celedoniana.

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International Rock Gardener

Number 132 The Scottish Rock Garden Club December 2020

Alan McMurtrie Iris ‘Storm’

- and good health in 2021!

We begin by profiling the late Carlos Celedón, who

fig.1: Carlos Celedón, the main protagonist, who

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Introduction and background

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fig.3: Valparaiso Region and the two distinct known locations of

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fig.7: Alstroemeria pulchra, scattered along the

fig.8: Yramea cytheris, the southern fritillary

Attractive, familiar Calceolaria polifolia [fig.9], a

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fig.12: Haploppapus velutinus is fairly common at mid-

We're now getting close to the point where the public

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fig.14: Although recorded from 7 regions up to 2600m,

Familiar, common and hummingbird-adapted

Of the two ferns we can say

fig.15: Adiantum chilense, one

fig.16: Another frequent pteridophyte in the Río

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fig.18:Tetraglochin alatum is one of the shrubby

fig.22: A small Cistanthe

fig.23: Cristaria dissecta, seen as a

That covers all we know about the relevant

Slightly lower down from the cistanthe

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fig.24: Cistanthe celedoniana habitat at the La Campana National Park.

fig.26: Malesherbia fasciculata, a very

At the north-exposed lower site (965 m)

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fig.31: Microphyes minima (Caryophyllaceae). This representative of a little-known genus looks

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Diurnal plant tropism

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fig.42: Cultivated individual transferred from

fig.43: Cistanthe celedoniana. A collected plant in

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fig.53: (a) Cistanthe celedoniana

fig.54: (b) Cistanthe celedoniana

fig.55: (c) Cistanthe celedoniana

fig.56: (d) Cistanthe celedoniana

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fig.58: (f) Cistanthe celedoniana

fig.59: (g) Cistanthe celedoniana

fig.60: (h) Cistanthe celedoniana

fig.61: (i) Cistanthe celedoniana