Bradleya 39/2021 45
A new species of Cumulopuntia (Cactaceae) from north-western
Paul Hoxey1,2 and Martin Lowry3,4
1 34 Stonehill Road, Great Shelford, Cambridge, CB22 5JL, UK (email: email@example.com).
2 Gibraltar Botanic Gardens, Red Sands Rd, GX11 1AA, Gibraltar.
3 Northgate Avenue, Macclesﬁeld, Cheshire, SK10 3AE, UK (email: firstname.lastname@example.org)
4. IOS Executive Board member (http://succulentresearch.org/)
Photographs: Paul Hoxey and Martin Lowry
Summary: A new species of Cumulopuntia (C.
flexibilispina) from north-west Bolivia, which
forms hemispherical clumps with dense flexible
white spination, is described and illustrated. The
species is only known from very high elevations
between 4,400m and 4,650m. The new species
is compared with and key differences are noted
from four other high elevation cumulopuntias: the
widespread C. boliviana (including C. boliviana
subsp. dactylifera) from Peru, Bolivia and Argen-
tina; C. glomerata (C. boliviana subsp. ignescens)
which is found at similar high elevations but in
more arid conditions further to the west in Peru,
Bolivia and Chile; C. chichensis from southern Bo-
livia and northern Argentina and finally C. echi-
nacea from southern Peru and northern Chile.
Zusammenfassung: Eine neue Cumulopuntia-Art
(C. flexibilispina) aus dem Nordwesten Boliviens,
die halbkugelförmige Gruppen mit dichter, flexi-
bler, weißer Bedornung bildet, wird beschrieben
und abgebildet. Die Art ist nur aus sehr hohen
Lagen zwischen 4,400 m und 4,650 m bekannt.
Die neue Art wird mit vier anderen Cumulopun-
tien hoher Lagen verglichen und die wesentlichen
Unterschiede zu diesen werden beschrieben: zu
der weit verbreiteten C. boliviana (einschließlich
C. boliviana subsp. dactylifera) aus Peru, Bolivien
und Argentinien, zu C. glomerata (C. boliviana
subsp. ignescens), die in ähnlich hohen Lagen,
aber unter trockeneren Bedingungen weiter west-
lich in Peru, Bolivien und Chile vorkommt, zu C.
chichensis aus Südbolivien und Nordargentinien
und schließlich zu C. echinacea aus Südperu und
During a visit to north-west Bolivia, south of the
town of Charazani in 2003, one of us (Martin Lowry)
encountered an opuntioid species growing at very
high elevations and later illustrated it (Lowry & Men-
doza 2011, Fig. 2). At the time this plant was identi-
fied as Cumulopuntia boliviana, a widespread taxon
from Bolivia, Argentina and Peru. There, at 4410m
above sea level, it was growing in the company of
two other high elevation cacti, Echinopsis maximili-
ana and Austrocylindropuntia floccosa.
Martin returned to north-west Bolivia with
the first author in 2017 and, when travelling
north from Charazani towards Ulla-Ulla across
a flat desolate plain, again at around 4,400m, a
number of hemispherical-shaped cacti were en-
countered. Martin identified them as C. boliviana
and remarked they appeared to be similar to the
plants he had found in 2003 south of Charazani,
about forty kilometres to the south. The plants
form hemispherical clumps to 50cm across and
about the same height and consist of many small
segments (Figure 1). The most striking feature of
the plants is the spination. Young segments have
brown spines (Figure 2) but these fade to a glassy
white at maturity and become very soft and flex-
ible (Figures 3–4). We carefully examined these
plants and came to the conclusion they had a num-
ber of characteristics that separated them from C.
boliviana as we understood the species. The soft
flexible spination in particular is unlike C. bolivi-
ana or related species. Continuing our exploration
of the region, we found the plants at several more
localities, including north of Ulla-Ulla where they
grow with Punotia lagopus, an association that we
have not encountered before with Cumulopun-
tia. At another locality we found plants growing
with Echinopsis maximiliana, another taxon well
known to grow at very high elevations. When re-
turning from Charazani to La Paz we stopped at
Martin’s original locality from 2003 (LM0433) and
collected a specimen which was later preserved
and deposited at LPB.
In all populations the characteristics of the
plants were consistent with little variation between
specimens, except for some minor differences in
the number and the flexibility of the spines. We
believe this plant is an undescribed species of Cu-
mulopuntia which we name here for the first time.
Fortunately, material was found in flower and with
ripe fruits from the previous year which allowed a
full description to be made in situ.
Cumulopuntia flexibilispina Hoxey & M. Lowry
Diagnosis: The species differs from all other
Cumulopuntia in forming hemispherical mounds
of numerous segments with dense and flexible
spination, with 15–20 spines per areole. Young
segments have distinctive brown spines which
turn glassy white and become flexible at maturity.
Type: BOLIVIA. Dept. La Paz, south of Charaza-
ni, S15°17’33.1” W069°02’30.9”, 4,500m, 8 Febru-
ary 2017, P. Hoxey and M. Lowry 005 (Holotype:
LPB). The type specimen is illustrated here prior
to preservation (Figure 5).
Etymology: The epithet is taken from the Latin
adjective ‘flexibilis’ meaning flexible combined
with the noun ‘spina’ for spine which refers to the
flexible nature of the spination.
Description: Plants form hemispherical
clumps to 50cm diameter and tall, with densely-
packed segments. Roots fibrous. Segment (Figure
6), ovoid, rounded, about twice as long as wide,
50–60mm long by 25–30mm wide, epidermis
green and smooth. Upper half of segment weak-
ly tuberculate. Areoles restricted to upper two
thirds of segments, 2–3mm high by 1–2mm wide
with white felt. Lower areoles spaced 10–15mm,
without spines but with numerous light yellow
glochids, 2mm long. Areoles near the apex of the
segment more closely spaced, 5–10mm apart, with
spines, glochids sometimes absent. Spines about
15–20, 15–30mm long, glassy white, somewhat
variable from almost hair-like, through to very flex-
ible bristle-like spines, often contorted or twisted
to slightly stronger, often straighter, but still flex-
ible spines. The latter sometimes with brown tips.
Figure 1. Typical hemispherical clumps of Cumulopun-
tia flexibilispina (PH1338.01 / LM1139.01) found east of
Figure 2. Cumulopuntia flexibilispina (PH1338.01 /
LM1139.01) with immature segments with brown spi-
nation surrounded by mature heads with white spines.
Figure 3. Typical flexible white spination of Cumu-
lopuntia flexibilispina (PH1338.01 / LM1139.01).
Figure 4. A particularity densely spined example of Cu-
mulopuntia flexibilispina (PH1339.01 / LM1140.01).
Bradleya 39/2021 47
Immature segments easily distinguished from ma-
ture segments with brown spination, noticeably
more tuberculate at the apex and with a small
green leaf, 1–2mm long, sometimes present at the
base of the areole. Flower (Figures 7–8) 30–35mm
long by 20–25mm wide. Flower tube 28mm long
by 16mm wide at top and approximate 5mm thick.
Green and completely naked in lower two thirds.
Areoles mainly concentrated around and near
the top of the flower tube; rarely found below.
Areoles approximately circular with white felt
and a small green scale. Between 10–15 flexible
spines to 10mm long, glassy white but occasion-
ally reddish-brown in upper parts. Ovary 4mm
wide across top and 5mm deep. Nectar chamber
3mm high. Style 15mm long and white. Stamens
numerous from the top of the nectar chamber to
the top of the tube, whitish, approximately 7mm
long, anthers small and light yellow. Sepals green,
4mm long by 2mm wide, pointed, slightly fleshy.
Petals yellow, in several overlapping rings, 12mm
long by 6mm wide, rounded top, about 20–25 in
total. Fruit (Figure 9) 35mm long by 20mm wide,
greenish yellow at a year old and easy removed.
Semi-circular depression in the top 15mm wide
and 10mm deep. A few small areoles on the rim,
less than 1mm diameter with about 6 white flex-
ible spines, almost hair-like, about 10mm long.
Seed cavity 15mm high by 6mm wide with 8
seeds. Seeds (Figure 10) pyriform, 5mm long by
4mm diameter, the funicular envelope slightly ru-
gose and off-white to pale brown in colour with a
prominent funicular girdle up to 0.5mm thick.
Habitat and Distribution: C. flexibilispina is
found growing between 4,390 and 4,650m, a re-
markably narrow elevation range. We encoun-
tered plants at five localities (outlined in Table 1)
but the first four listed can probably be considered
to belong to one fragmented population which
stretches from south of Charazani northwards to
Laguna Antaquilla, for approximately fifty kilome-
tres. Plants were not observed continuously over
Figure 5. The type specimen of Cumulopuntia flexibi-
lispina (PH1355.03 / LM0433.02) prior to preservation.
Figure 6. Segment of Cumulopuntia flexibilispina
(PH1355.03 / LM0433.02).
that distance and appear to be divided into small
discrete populations. The fifth locality is signifi-
cantly disjunct from the others being 300km away
which suggests C. flexibilispina may be found in
high elevation habitats in the intermediate region.
The type locality is at 4,500m where specimens
of C. flexibilispina are found on heavily grazed
hillsides growing with Austrocylindropuntia floc-
cosa and Echinopsis maximiliana. The plants can
be found on rocky outcrops or on grassy slopes.
The two populations near Ulla-Ulla grow on a flat
plain in stony ground with little other vegetation.
There are scattered clumps of grasses and a few
low bushes but C. flexibilispina is the most visible
plant in this inhospitable habitat. At the popula-
tion north of Ulla-Ulla, C. flexibilispina grows to-
gether with Punotia lagopus (Figure 11). This is an
interesting association and, as far as we are aware,
the first time the genera Punotia and Cumulopun-
tia have been found growing sympatrically. At this
place we also observed that the flowers had been
infested with small red mites (Figure 12).
The most northerly population encountered
is North of Laguna Antaquilla on the road to Pel-
echuco. Here the plants grow at 4,650m (the high-
est elevation we know to date) on gentle slopes
with rocky outcrops (Figure 13). The area is grazed
but a little grass and some alpines can be found.
The animals appear to prefer to graze the water
logged areas nearer the lake with green grasses.
The nearby rocks are home to Echinopsis maximil-
iana. This is a remarkable elevation to find cacti,
broadly equivalent to the maximum recorded for
Austrocylindropuntia floccosa and Punotia lagopus
and only C. glomerata (C. boliviana subsp. ignes-
cens) is confirmed to grow significantly higher at
4,830m (Hoxey, 2016). Together these four locali-
ties make up the ‘core’ population.
A fifth and outlying population was encoun-
tered on the road over the mountains from Pataca-
maya on the highway Ruta National 4 to Luribay
south of La Paz where we found a few plants at
4500m. The habitat was not grazed and as a result
the slopes were covered with tufts of ichu grasses.
The plants of C. flexibilispina were rather well
hidden in the grasses so not easy to see without
searching. The plants are a little different from
the ‘core’ populations forming clumps a little wid-
er than tall and with spination that is generally
straighter and stiffer although still relatively flex-
ible. The spination is still glassy white, however,
and the immature spines are a red-brown colour
consistent with the other C. flexibilispina popula-
The area of distribution is limited by elevation
but furthermore the plants appear further restrict-
ed to fragmented populations within the general
area. The precise reason why the species appears
unable to grow at lower elevations is unknown.
We saw no evidence of asexual propagation. The
segments are firmly attached to the mother plant
and are unlikely to be knocked off and dispersed
to form new plants so reproduction is probably
mainly by seed. At the type locality we were for-
tunate to find a couple of small seedlings and one
is illustrated in Figure 14.
C. flexibilispina is presently only known from
Bolivia and restricted to the Department of La
Paz. The populations near Ulla-Ulla are extremely
close to the border with Peru so it is highly likely
the species will be found there. So far, we have
not had the opportunity to confirm this.
Conservation Status: It is only possible to as-
sign a provisional conservation rating to this spe-
cies as at present our knowledge of the distribution
is incomplete. Based on a core population with an
extent of occurrence estimated at 500km2 and a
significantly smaller area of occupancy with only
four known localities a conservative IUCN (2001)
classification of Endangered (EN) seems appropri-
ate. However, if further populations are found be-
tween the core area and the disjunct locality near
Locality Lat. / Long. Elevation Associated
4km east of Ulla-Ulla S15°04’09.5” W069°14’00.8” 4400m
5km north of Ulla-Ulla S15°01’02.0” W069°16’23.9” 4390m PL
North of Laguna Antaquilla
on road to Pelechuco
S14°49’32.5” W069°13’53.4” 4650m LM
South of Charazani* S15°17’33.1” W069°02’30.9” 4510m AF LM
west of Luribay, on road
S17°12’57.5” W067°48’28.4” 4520m
Table 1. Overview of Cumulopuntia flexibilispina habitats (AF= Austrocylindropuntia floccosa LM=Echinopsis
maximiliana, PL=Punotia lagopus, *=type locality)
Bradleya 39/2021 49
Figures 7–14: 7. Cumulopuntia flexibilispina in flower
(PH1338.01 / LM1139.01). 8. Flower section Cumu-
lopuntia flexibilispina (PH1355.03 / LM0433.02). 9.
Fruit of Cumulopuntia flexibilispina (PH1338.01 /
LM1139.01), smaller and with fewer seeds than is typical
for other high elevation Cumulopuntia. 10. Seeds of Cu-
mulopuntia flexibilispina (PH1339.01 / LM1140.01). 11.
Cumulopuntia flexibilispina (PH1339.01 / LM1140.01)
growing with Punotia lagopus (PH1339.02) near Ulla-
Ulla. 12. Cumulopuntia flexibilispina (PH1339.01 /
LM1140.01) with red mites in the flower. 13. A large
clump of Cumulopuntia flexibilispina (PH1344.02 /
LM1145.01) growing at 4,650m with many fresh seg-
ments. 14. A small seedling of Cumulopuntia flexibi-
lispina (PH1355.03 / LM0433.02).
Figures 15–16. Cumulopuntia flexibilispina (PH1338.01/ LM1139.01) East of Ulla-Ulla, La Paz, Bolivia, 4,400m.
Figures 17–18. Cumulopuntia glomerata (PH1267.01) Volcan Taapaca, northern Chile, 4,800m.
Figures 19–20. Cumulopuntia echinacea (PH1202.06) Putre, northern Chile, 3,700m.
Figures 21–22. Cumulopuntia boliviana (PH13785.01 / LM1170.01) South-west of Viacha, La Paz, Bolivia,
Bradleya 39/2021 51
Luribay there will be grounds for downgrading
the classification. There are no immediate threats
to the species as the land at such high elevations
is only suitable for grazing, an activity that has oc-
curred there for centuries.
Discussion: Cumulopuntia flexibilispina joins
a select group of four cactus species: Austrocylin-
dropuntia floccosa, C. glomerata (= C. boliviana
subsp. ignescens), Echinopsis maximiliana and Pu-
notia lagopus which are found at very high eleva-
tions above 4,500m (Hoxey, 2016). C. flexibilispina,
together with P. lagopus, are the only two species
which are restricted to these elevations. P. lagopus
has a range of 4,300–4,700m (personal observa-
tions) although there are unconfirmed reports that
it can grow higher. C. flexibilispina has a range of
4,390–4,650m which, by nearly 100m, takes the re-
cord for the cactus species with the highest mini-
mum elevation known at 4390m.
There are several other taxa which need to
be compared with C. flexibilispina. Here we in-
clude other high Andean Cumulopuntia: C. bo-
liviana (including subspecies) and C. chichensis.
For each taxon we include two illustrations, one
of a complete plant and another of a close up of
the segments to facilitate comparison between
them (Figures 15–26). The very widespread spe-
cies C. boliviana was considered to consist of four
subspecies (subsp. boliviana, subsp. dactylifera,
subsp. ignescens and subsp. echinacea) by Hunt
(2002). A molecular study (Ritz et al., 2011) does
not support this classification because only C. bo-
liviana subsp. dactylifera has a close relationship
to C. boliviana subsp. boliviana. More recently,
Hunt (2016) has suggested C. boliviana subsp. ig-
nescens should be treated as a good species and
we agree. Hoxey (2020) has also shown that the
correct name for this taxon at species level within
the genus Cumulopuntia is C. glomerata (Pfeiff.)
Hoxey (Basionym: Pereskia glomerata Pfeiff.)
which must not be confused with Maihueniopsis
glomerata (Haw.) R.Kiesling (Basionym: Opuntia
glomerata Haw.) which is a completely different
species. It should also be noted that the epithet
‘ignescens’ still has priority at the level of subspe-
cies. We also consider C. echinacea as a good spe-
cies in this paper.
The hemispherical shape and densely packed
segments of C. flexibilispina (Figures 15–16) are
most reminiscent of C. glomerata (Figures 17–18)
Figures 23–24. Cumulopuntia boliviana (subsp. dactylifera) (PH1002.03) South of Putina, Puno, Peru, 4,010m.
Figures 25–26. Cumulopuntia chichensis (LM1237.01) Yunchara, Tarija, Bolivia, 3,640m.
C. flexibilispina C. boliviana (incl.
C. chichensis C. glomerata (=
C. boliviana subsp.
clumps, 50cm wide
clumps, to 50cm+
across but only 1 or
2 segments tall.
or low spreading
clumps to 1m ×
clumps, to 50cm+
across, usually a
little shorter than
clumps of many
60cm across and
Segments 50–60 × 25–30mm.
Upper half of
Lower third of
third with areoles,
third with closely
spaced areoles with
spines but glochids
40–60 × 25–30mm.
Lower half of
half of segment
and with areoles.
Upper areoles with
spines, lower with
75–95mm tall by
diameter on upper
half, smooth below.
50mm long by
green. Lower half
areoles, upper half
with areoles that
40–90 × 45–55mm.
A small number of
large felted areoles
on the upper half
of the segment. No
Spines 15–20, 15–30mm
from hair-like, or
or twisted, to
but still flexible
yellow to yellow-
12–16, straight, but
spine to 50mm
long, weaker spines
mm long, reddish,
fading to light
acicular but a little
30–120mm but to
strong and thick,
reddish brown to
Flower 30–35 × 20–25mm.
Flower tube 28 ×
16mm, green and
in lower two thirds.
the rim, 10–15
flexible spines to
10mm long, glassy
white but some
Petals yellow or
tube 30 × 30mm,
areoles only on rim
with 5–8 spines to
10mm long, white.
yellow. Flower tube
areoles near rim
with bristly spines
or with glochids.
60mm long by
50mm wide, petals
red or orange-red.
Flower tube 30 ×
green, areoles only
on rim with 10–15
spines to 15mm
35–55 × 35–55mm
wide, petals yellow.
Flower tube 12 ×
green, areoles only
on rim with about
10 spines 20mm
Fruit 35 × 20mm wide,
greenish yellow. A
few small areoles
on the rim with
about 6 white
about 10mm long.
except for a few on
the rim. Generally
50 × 40mm,
mainly on rim with
glochids and about
12 spines 5–15mm
60mm long by
Areoles on rim
with about 12–15
spines to 60mm
long, straight but
25–40 × 25–30mm.
Areoles on rim
with about 4–10
spines to 15–60mm
4,390–4,650m 3,530–4,280m 3,090–3,940m 3,370–4,830m 2,960–4,190m
Table 2. Summary of the principal characters of C. flexibilispina compared with C. boliviana (from habtat near
La Paz), C. glomerata and C. chichensis and C. echinacea. Data for C. flexibilispina, boliviana and glomerata from
authors’ original observations. Data for C. chichensis and echinacea from orginal descriptions.
Bradleya 39/2021 53
which grows further to the west and south in gen-
erally drier situations but is also a plant of very
high elevations. The basic plant morphology of
both species is almost certainly an adaptation to
the cold of high elevations. Neither species has de-
veloped a dense covering of wool or hair although
the spination of C. flexibilispina does show adapta-
tions in that direction. The two species can be eas-
ily distinguished with C. glomerata having much
stronger and reddish or yellow-red spination, red
or orange-red flowers and fruits with long flexible
spination on the rim.
The other two Opuntioids which grow at very
high elevations (Austrocylindropuntia floccosa and
Punotia lagopus) have evolved the ability to cover
themselves in dense hairs. Unlike C. flexibilispina
and C. glomerata they tend to form clusters of stems
with a flatter profile, wider than tall. Perhaps the
wool allows them to do this whereas C. flexibilispi-
na and C. glomerata need to form the hemispheri-
cal shape to retain heat more effectively without
the ability to insulate themselves with hair.
Cumulopuntia echinacea (Figures 19–20) is of
rather limited distribution in southern Peru and
northern Chile and has extremely strong spines,
perhaps the longest in the subfamily Opuntioideae,
reaching 15cm long. One of us (Hoxey) has found
C. glomerata and C. echinacea growing sympatri-
cally in southern Peru and we also consider this a
good species and easily distinguished from C. bo-
liviana as well as C. flexibilispina due to the flatter
growth of the clumps, the larger but less densely
packed segments and the very robust and longer
C. boliviana (subsp. boliviana) is a very wide-
spread species with variable spination and almost
spineless on occasion. For comparison with C. flexi-
bilispina we use plants found south and south west
of La Paz because they are populations that are rea-
sonably close by and representative of this species.
These populations of C. boliviana (Figures 21–22)
form clusters of stems which are much wider than
tall, usually only reaching a height of two or three
segments above ground level, although plants in
other populations can grow into a more hemispher-
ical shape. The spination is variable in length but
generally consists of 5–8 spines, 4–6cm long which
emerge from the areoles on the upper part of each
segment. The spination is usually yellow but can
fade to a dark grey with age. Flowers of C. bolivi-
ana are similar to C. flexibilispina but a little larger
and are usually yellow but can be orange in some
populations. C. flexibilispina is easily distinguished
from C. boliviana due to its white flexible spination
and a much higher spine count. The flowers of C.
flexibilispina are also exclusively yellow.
C. boliviana subsp. dactylifera is a northwards
extension of C. boliviana into southern Peru and
plants are found to the west and north of Lake Titi-
caca and north-west of there. The main differences
regarding C. boliviana are the spination which is
generally weaker although variable, and the more
elongated segments. We are not convinced that
this subspecies is worthy of recognition and in this
paper include it in our concept of C. boliviana. It
does not appear to be geographically isolated from
C. boliviana (subsp. boliviana) nor are the charac-
ters sufficiently distinct. However, we illustrate a
plant (Figures 23–24) growing about forty kilome-
tres from Azángaro, the type locality of Opuntia
dactylifera, the basionym of C. boliviana subsp.
dactylifera which is representative of this taxon. It
is also about fifty-five kilometres from the nearest
populations of C. flexibilispina.
Cumulopuntia chichensis from southern Bolivia
has many characteristics similar to C. boliviana but
it has white spines which are more robust, longer
and often contorted (Figures 25–26). Interestingly,
it has brown spines on young immature segments
which turn white at maturity like C. flexibilispina
but the spines of C. chichensis are less numerous
and much stronger. It is possible that C. chichensis
is most closely related to C. echinacea which re-
places it to the west.
Table 2 compares characteristics of C. flexibi-
lispina with the other high elevation Cumulopuntia
We are especially gratefully to Dra Carla Maldona-
do, director of the Herbario Nacional de Bolivia,
and to Alfredo F. Fuentes Claros of the same institu-
tion, for providing us with permits to access and
collect within the Madidi National Park and Apolo-
bamba Natural Integrated Management Area.
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