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Succulent plants from down under Adventive Plants (Part 5):

Cakile maritima ssp. maritima - Aloe maculata - Opuntia monacantha

XI. Cakile maritima ssp. maritima

I have to admit that Cakile maritima Scopoli 1772 ssp. maritima was kind of mystery plant for me for
few months, after I have noticed it on some Coromandel Peninsula beaches like the legendary Opoutere
beach and Whangamata beach, growing along with native sand dunes grasses or alien succulent plants.
My first thought was that it has to be a native plant and I was looking for months unfortunately in this
direction, but not getting even a clue of what it might be. I even have in Part 3 of this series o photo
showing Carpobrotus edulis growing amongst native succulent plants that native succulent was Cakile
maritima (1). One day I posted a short message and links to some photos on Didge Rowes Xerophytes
NZ notice board and I finally got the answer someone new in an instant whats all about.

1. Cakile maritime ssp. maritima growing with Carpobrotus
edulis at Whangamata, Coromandel. By the time this photo
was taken I was pretty sure that this must be a native plant.
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There are quite a few things worth knowing about this plant, dull and insignificant at first sight, but full
of surprises on a closer look. First of all the well known Searocket by its vernacular name is a highly
resistant halophyte plant; it takes it to the extreme being able to survive concentrations of over 400mM
of NaCl (salt). I have briefly mentioned in a previous article (2) how halophyte plants manage to survive
killer concentrations of salt, but Cakile maritima is somehow different from Sarcocornia quinqueflora.
By being annual plants (3), with fruits ripen in summer shortly after the flowering period, and seeds
germinating in winter, the seeds are therefore very tolerant to higher salt concentrations, like most of the
annual halophytes. Not all seeds are germinating at the same time; this would be very risky and unwise
in adverse environments as the fore dunes usually are - some of them form long lasting seed banks
ensuring a constant germination over a period of time, ensuring new generations no matter what, if by
some extreme conditions entire populations or seedlings became extinct. It is this asynchronous
germination of most annuals in hazardous environments that maintain the continuity of the species. As
expected the seeds have also a high longevity, it maintains for three years a high germination rate, being
able to germinate on occasion even after many years. Seed banks are very resistant to soil disturbances
even if these occur several times a year for two or three years in a row and this may happen in very
unsettled sand dunes habitats. This is actually the preferred habitat, not the wet. It is very resistant to salt
spray and transient sea water inundation; capsules are floating on sea water and retain the germination
capability of the seeds, although these are reluctant to germinate in extremely high salinity environments
dispersal by sea (waves, currents, winds) seem to be a very efficient vector.

Although beach and dune sand is (beside its high salt content) a very poor substrate, with barely any
nutrients with nitrogen content, Cakile maritima has a very positive reaction to local and / or casual
nitrogen enrichment showing explosive growth and vegetative development, unlike most of the typical
plants sharing this habitat (like Spiniflex spp., Calystegia spp., Disphyma spp., Carpobrotus spp., etc.
showing poor growth or simply avoiding such areas), especially in association with mineralization.
Growth is also highly stimulated by perturbation events like persistent strong winds, burial by blown
sand, temporary sea water inundation, various soil disturbances, etc., drifting plants or plant parts
removed from their original locations being able to form extremely quickly the nuclei of early
succession fore dunes vegetation elsewhere.
2. Cakile maritima ssp.
maritima tussocks
scattered on the beach
at Whangamata.
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Cakile maritima ssp. maritima (belonging to Brassicaceae, the mustard family), is a low growing bushy
plant, with stems reaching only 10 30 cm, much branching from the base, stout and glabrous, from
sub-erect to decumbent, spreading and forming small mounds, usually 30 x 30 cm. Both stems and
leaves are succulent. The fleshy leaves form a very loose rosette and are narrow-oblong to ovate,
shallowly to deeply crenate, petiolate, 40 60 mm x 10 20 mm, with leaf margins usually entire but on
occasion shallowly crenate or sinuate. Stem leaves are usually much smaller, crenate, and usually having
a different shape from oblong to linear to linear-spathulate. The four petal flowers are hermaphrodite
have 10 mm diameter and are whitish to pale lilac, growing in loose clusters on 1 3 mm long and
spreading pedicels, and pollinated by bees, flies, beetles and moths. The inflorescence is straight,
thickening and elongating at fruiting. As the specific epithet shows the plant is fully confined to
maritime strandlines growing usually on sand and shingle and on the nearby fore dunes. Cakile maritima
is found around the shores of Western and Southern Europe and North Africa, several subspecies are
known showing significant differences in fruit morphology and leaf form (4). Another closely related
species - Cakile edentula is native to the east coast of North America. Both species have been
introduced on the Pacific coast of North America in some of the Pacific Islands, Australia and New
Zealand posing a moderate threat to native coastal flora. The primary danger is not the direct
3. Cakile maritima ssp. maritima at Whangamata.
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competition with native plants for resources, Cakile maritima is rather a pioneer plant, establishing
quickly in disturbed habitats, but not performing too well in dense vegetation areas; it has been
established that the plants from scarcely vegetated beaches are the main seed source for new plants in
grassy former fore dunes, the germination rate is so poor in such areas that an external seed flux is really
necessary to maintain the populations on the long run. It is therefore less effective and less threatening in
my opinion for the New Zealand coastal ecosystems than you might be considering in first instance. As
in its original habitat the importance of Cakile maritima in the conservation of the coastal habitats has
been acknowledged, I think that being a first succession pioneer plant, by dominating the drift vegetation
type and giving up in established environments, it is acceptable for New Zealand for conservation
purposes in highly degraded and volatile coastal ecosystems. It is apparently more of a dune initiating
plant, rather than dune building, and therefore preparing the environment for the secondary successions.
Cakile maritima has usually a higher salt tolerance than the secondary succession plants.

It is very interesting that the genus Cakile has a huge dispersal, an almost worldwide range. It is
considered that an ancestral Cakile sp. was originally a desert plant, being probably similar in habit to
Cakile arabica. From some reason this ancestral Cakile made the transition from a desert to a maritime
4. Cakile maritima ssp. maritima at Opoutere, Coromandel.
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habitat and experienced a consequent increase in global range by using the waterways for medium and
long-range dispersal. As the salty seawater does not damage the floating capsules of the plant or the
seeds themselves for quite a time it might be a reasonable explanation (5). To complete its profile even
if a warmth and sun loving plant (does not grow in shade at all) it is also frost hardy during winter,
surprisingly for a coastal plant, possibly due to recessive genes inherited from the ancestral desert

Very surprisingly the plant used to be cultivated in Great Britain not only for its aspect (very interesting
indeed, but not really showy for my taste) but also for culinary uses; rich in Vitamin C but very bitter,
the plant (pretty much everything leaves, stems, flower buds and immature seedpods) was used mainly
as a flavouring spice, raw or cooked.

The plant is quite wide spread in New Zealand (it has been naturalized in 1940), but only in the North
Island: Kaitaia, Bay of Islands, Hukerenui (Northland), Whangapararoa (Auckland), Thames, Opoutere,
5. Although both are beaches with eastern to northeastern exposure at the Pacific Ocean,
the Opoutere plants have a somewhat different look, smaller sized, showing clearly the
pressure of a very harsh environment. In contrast, the Whangamata plants have that
peaceful nice look of cultivated plants. This plant was photographed at Opoutere.
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Whangamata (Coromandel Peninsula), Bay of Plenty, East Cape, Gisborne, Hawke's Bay; Waipawa,
New Plymouth, and Wellington. I have seen the plant in two of the locations mentioned in Flora of New
Zealand, Vol. 4 Opoutere and Whangamata (Coromandel Peninsula), abundantly but never causing
high-density infestations, on sandy beaches close to the high tide line.

XII. Aloe maculata

I have already briefly mentioned Aloe maculata (Aiton) Haworth 1804 (usually referred as Aloe
saponaria (Aiton) Haworth 1804 by New Zealand botanists) in relation with the succulent plants of
Rangitoto Island, Hauraki Gulf, but I think that going a bit into details will have some relevance as this
6. 8. Taking a
picture of Cakile
maritima ssp.
maritima at Opoutere
(above left) and
habitat pictures at
Opoutere (above right
and left).
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plant even if extremely common and very adequate for beginners seem to be everyones favourite
and often encountered even in fancy collections.

It is usually quite tricky to identify the spotted Aloe plants, some of them extremely similar when not in
flower, but Aloe maculata is fairly distinctive (6). It forms clumps of stemless (or forming in time a very
short stem) succulent rosettes usually to 25 30 cm high (on occasion up to 50 cm high) and 25 - 30 cm
wide (on occasion 40 50 cm wide). The succulent leaves usually 12 20 in a rosette may vary in
shape (7) but are usually elongated triangular broader at the base and getting thinner toward the tips; in
some plants the leaves may have an almost belt-like appearance. The green leaves are copiously spotted
with whitish to light green spots on the upper surface, arranged in vaguely transversal stripes, more or
less evident, sometimes in wavy designs. Sharp brownish marginal teeth are present. The upper side of
the leaves is usually plane to slightly concave while the lower side is in most of the plants spotless and
convex. The plant is very prolific in sending rhizomes in all directions to shoot new plantlets eventually
forming large clumps. The axillary inflorescence is branched and bears usually red to red-orange
flowers, but there are also salmon-red, orange and yellow flowering forms, and of course all possible
9. Aloe maculata, young plants growing directly on lava block on Rangitoto
Island, Hauraki Gulf. Collected debris seems to be the only moisture retainer.
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nuances in-between. Aloe maculata appears to have (at least this is my experience) a limited self-
compatibility on occasion otherwise it is self-sterile.

Aloe maculata has a wide distribution in South Africa (reaching from the Cape Peninsula to the Western
and Eastern Cape Provinces through Orange Free State and KwaZulu-Natal and Mpumalanga), Lesotho
and Zimbabwe in a variety of habitats from grassland to thicket and rocky outcrops. It occurs mostly in
milder coastal climates, with humid winters, but also deep into the inland at higher altitudes and in
summer rain areas. Having such a wide dispersal area it is not threatened at all, but present in large
numbers in numerous locations.

10. This is the main rosette
of the plantlet collected
from Wenderholm in July
2003. Although I have
tried to grow it hardy and in
full sun there is nothing like
a wild plant.
11. Typical inflorescence
of Aloe maculata the
flat raceme.
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Aloe maculata is definitely a very successful plant, not only for being a favourite landscaping plant, but
also for becoming if not naturalized then at least a casual and persistent presence, a typical garden
escape, basically in all tropical and subtropical regions of the world, especially in the Mediterranean
region, India, China, Australia, New Zealand, some of the Pacific Islands, United States (especially in
the south-west where numerous escapes have been observed) and even South America.

This plant is extremely popular here in New Zealand to begin with, probably the most cultivated Aloe.
The easy-going nature of this plant and the beautiful foliage and flowers contributed to its early success
in gardens, escaping probably very soon in nature. The first New Zealand collection is dated 1883 (one
of the first succulent plants naturalized here) and was made by T. F. Cheeseman in Remuera (Auckland),
identified as Aloe latifolia (Haworth) Haworth 1804, now considered to be a synonym.

Aloe maculata is quite widespread here in New Zealand being one of the very few fully naturalized
succulent plants (opposed to adventive or casual appearances) and there is no wonder that it managed to
do this, as it is very popular in cultivation. It is a therefore typical garden escape, discarded garden waste
12. Aloe maculata a wild beauty from Rangitoto Island, Hauraki Gulf.
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being probably the most likely mean of dispersal. But unlike other escapes this plant has really managed
to form self-maintained populations occurring in several sites throughout New Zealand. The plant is
usually referred as Aloe saponaria (Aiton) Haworth 1804 by most New Zealand authors and although
the name is incorrect it looks like this tradition is hard to break.

Few significant wild populations have managed to establish in time mainly in the northern half of the
North Island - such as Mt Puheke in Karikari Peninsula and Tokerau Beach - Paihia (Northland),
Auckland City, Rangitoto Island (Hauraki Gulf) but also few South Island populations are scattered on
the coastal areas of Canterbury and Christchurch; as you can see it is confined to coastal areas.

I have seen this plant in nature in two different locations. The first time in July 2003 at Wenderholm, 40
km north of Auckland, in a Regional Park (which is actually a mixture of recreational area and nature
reserve) in a vegetation control area, at 100 m or so far from the waterline, in a light wood area. There
was a nice clump of several dozens of rosettes, some bigger mature plants, some just few centimeters
wide plantlets, forming a semi-circular colony around the base of a big tree. It had some sun there, but
13. Same plant as above. In Rangitoto I have seen just isolated loose
groups of plants, few dead ones too, scattered from place to place on
the eastern shore, but not large clumps as in Wenderholm.
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probably mostly placed in shade, the plants looked more like cultivated plants, very nice however. It was
a small but well settled population. I took a small rosette, a plantlet of ca. 5 cm diameter, which became
the starting point of vegetative reproduction in my greenhouse and rock garden, giving away some other
clones as well. Unfortunately I didnt manage to take pictures of this nice clump (I didnt have a digital
camera at that time), and one year later I was surprised to see that this lovely group of plants has been
destroyed and replaced with native vegetation, most likely the normal outcome for any intruder in a
controlled vegetation area. They must have done a pretty good job, as I havent seen re-growth of the
Aloe maculata since.

The second location is Rangitoto Island (Hauraki Gulf), where I have seen few sensational plants during
my second trip there in January 2008. I wrote about the Rangitoto plants in Part 4 of this series; these are
the plants that have impressed me most, truly wild plants growing on bare lava fields covered with
lichens, having mostly debris around them as moisture retainers, just few of them finding shallow rocky
substrate but no organic matter at all. All plants were displaying the brownish signs of stress inducted
possibly by lack of water or excessive sun exposure, or by both - these were probably the most beautiful
14. The young plantlets from Rangitoto seem to be according to the red-
brown colours displayed quite stressed by the scarcity of the water, or by
the intense sun exposure, or by both, but this is life in the wild.
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succulent plants I have seen in the wild. These plants have, I like to underline this, real character. Not
fancy chaps, but true survivors.

XIII. Opuntia monacantha

Opuntia monacantha Haworth 1819 is probably the most common opuntioid, everyone seem to know it,
have seen or had one, and so on. There is nothing special about this plant, widespread and almost a pest
even in its own habitat, but definitely there was a pleasant surprise for me as I saw it in nature (by pure
chance) on a sunny day spent with my family at Eastern Beach, Howick, Auckland.

No surprise for a very common cactus there is some confusion around it. Adrian Haworth described
the species in 1812, in Synopsis Plantarum Succulentorum. Haworth based his description on a plant
from Barbados, outside the natural distribution area. It is known today that it was introduced there and
that the only native Opuntia is Opuntia dillenii. He also refers also (with a question mark) Cactus
monacanthos Willdenow 1814 as synonym, but this indication of doubt makes under the current rules
impossible to consider Willdenows a basionym for that of Haworth, and therefore the quite often cited
name Opuntia monacantha (Willdenow) Haworth 1819 is not correct. Also the name Opuntia vulgaris
of different authors, including Miller, has been misapplied quite often by Australian and New Zealand
authors; due to this confusion in early New Zealand botanical records the plant is referred as Opuntia
ficus-indica, the source of a chain of confusion for many decades. Opuntia vulgaris Miller has now been
typified to become a synonym of Opuntia ficus-indica (Linnaeus) Miller.

The plant has usually a tree-like appearance, being usually 1.3 4.0 m tall, on occasion even more, and
having a terete trunk; in some plants this trunk is absent, the plants having then a rather shrubby
appearance. The pads are glossy green, obovate to oblong, narrowed at the base, usually 10 30 cm long
15. Opuntia monacantha,
the bigger plant seen at
Eastern Beach, Howick,
Auckland. It grows on
almost vertical cliffs
between grassy patches.
From place to place
organic matter seems to
have been accumulated in
the cracks, some moisture
is available as well from
the ground water.
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and 7.5 12.5 cm wide, with large areoles of 3 5 mm diameter and bear brownish 2 3 mm long
glochides. The spines are quite sparse on the joints, usually 1 or 2 per areole, sometimes 3 or even
absent, but densely on the main trunk up to 12 per areole. The spines are erect or spreading, usually
grayish with dark brown tips. The leaves are conic and 2 4 mm long, deciduous, not lasting too long.
The flowers have 5.0 7.5 mm diameter, the sepals being yellow with reddish midrib, while the petals
usually yellow, sometimes to orange. The fruit is obovoid with slightly depressed umbilicus and bears
light tan irregularly elliptic seeds of 3 4 mm. It usually flowers from mid spring to mid summer
provided strong sun and heat is present.

Opuntia monacantha is native to Argentina, Brazil, Paraguay and Uruguay, but has been widely
introduced and naturalized in almost all tropical and subtropical regions, from Florida to tropical Africa,
to South Africa, southeast Asia, Australia, New Zealand and some of the Pacific Islands, and I think
there must be some other places too. The worst infestations have occurred in Australia and South Africa.
It is also worthwhile mentioning that Opuntia monacantha was first recorded in China as early as 1625.

16. This is the smaller plant growing almost hidden by grasses some 1.0 1.5 above the
bigger plant.
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Opuntia monacantha (usually referred as Opuntia vulgaris by New Zealand authors, unfortunately a
compromised name, often misapplied to Opuntia ficus-indica) is nowadays not too wide spread here in
New Zealand in nature and this is quite a surprise considering the high occurrence in the past and the
Australian disastrous Opuntia introductions (8). However, it is the only fully naturalized Cactaceae in
New Zealand. Very popular in cultivation at some stage, Opuntia monacantha has managed to escape in
nature the most likely infestations being from abandoned plantations and dumped garden waste. At some
stage I think this was the fate of many garden plants going out of fashion and in older times no-one put
too much thought in how to discard the garden waste. I was told that the sand dunes of Mt. Maunganui
were a preferred place for such organic dumps for locals and few decades ago you had to consider very
wisely walking barefooted through the area as rooted Opuntia pads were littering the place. Reportedly
none has survived, but youll never know for sure unless you go there and browse thoroughly the area
(9). This fierce and invasive aggressor (considering the disastrous Australian events) has barely managed
to naturalize in New Zealand.

I think there is enough potential for isolated plants or small groups of plants to survive for extended
periods though. Opuntia monacantha can be on occasion very prolific in vegetative dispersal as pads
fallen on the ground root easily; also seed dispersal has been observed which is even worse. It just
17. Close up a pad from the bigger plant and the remainders of a flower.
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doesnt seem to be very effective in dense vegetation areas. It prefers warm and sunny banks and slopes,
sand dunes, beaches and rocky outcrops where it can form small self-maintaining populations, but it also
occurs in somewhat cooler locations. It is quite tolerant to marine exposure and also to moderate salt
concentrations. The New Zealand botanical literature refers following populations: Hokianga Harbour
and Mangonui (Auckland), Bay View - Napier (Hawkes Bay) (10), Awato (Nelson), Teviotdale Station,
Conway Flat, Waipara and near Waipara on the road to Weka Pass (Canterbury), Halswell Quarry
(Christchurch) (11) and - with a single known collection - Lowburn (Central Otago), which is the only
truly inland location, all others being coastal or near-coastal populations.

I have seen Opuntia monacantha in just one location Eastern Beach (Howick, Auckland), a location
which is not mentioned anywhere else so far I know, on the cliffs at the southern end of the beach,
between grassy patches. Unfortunately I didnt have the binoculars with me that day so that I couldnt
scan thoroughly the area. The two plants, hanging on almost vertical cliffs with eastern aspect were
probably at 3 4 m above the high tide mark, in a place, which can be accessed only at low tide. It could
have been more of them but I couldnt see them. As on the plateau, some 25 30 m above, there is a
residential area and many decades back, when Howick used to be well outside the settled Auckland
urban area, holiday baches were scattered, it looks like these plants are also remainders of dumped
garden waste. Looking at the residential properties above and the associated modern landscaping designs
this must have been the case. I think that I am not too far off maintaining that someone simply uprooted
and threw down the slope that big ugly cactus while builders were busy with the setup of the new
swimming pool. On the northern slope of the cliffs, exactly where the beach begins, in a more densely
vegetated area, there is also one of the most extensive Tropaeolum majus infestations I have seen in New

18. Eastern
Beach, Howick,
Auckland the
steep where I
saw the two
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I have seen just two plants, a mature plant having already flowered and, just 1 1.5 m above, a much
younger plant. This is rather strange that the much younger plant is placed above the older one. It is
virtually impossible to originate from a pad removed from the plant below and lifted somehow up by
strong wing, or even from seed dispersed upwards, there must have been some other plants higher on the
cliff, possibly hidden between grasses and shrubs, which I couldnt see. This younger plant originates
probably from seeds or pads dispersed from other plants above. I also discard a continuing source of
infestation form the plateau, this is highly unlikely.

I will probably return at some stage at Eastern Beach (making sure that I have the binoculars with me
this time) and I am pretty confident that new plants will be sighted. However, the unexpected finding of
these quasi-wild growing cacti was enjoyable enough.

19. From place to place you can see moist patches due to the water
sipping between the sediment layers from the ground water.
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Additional References:

A. G. Davy & al. - Biological flora of the British Isles: Cakile maritima Scop (Journal of Ecology, Vol. 94, 2006);

Flora of China (;

John J. Knapp, Joseph M. DiTomaso & al. Cakile maritima Scop. Plant assessment Form electronic version February 2003 (for the California
Exotic Pest Plant Council and the Southwest Vegetation Management Association);

Wided Megdice & al. Salt tolerance of the annual halophyte Cakile maritima as affected by the provenance and the development stage. (Acta Physiol.
Plant., vol. 29, 2007 published online by the Polish Academy of Sciences, Krakow 2007);

Bridget Ozanne & Charles David - Interesting Plants of Guernsey (not dated),;

Plants for a Future -;

Ben-Erik van Wyk & Gideon Smith - Guide to the Aloes of South Africa (1996);

E. Zimer Aloe maculata ( Cactus Romania Enciclopedie, 2006).

Further Readings:

R.W.G. Carter & Bill Carter - Coastal Environments An Introduction to the Physical, Ecological and Cultural Systems of Coastlines (1988);

John R. Packham, Arthur John Willis Ecology of Dunes, Salt Marsh and Shingle (1997);

My Notes:

(1) I will keep this error, as this series is not only an account on the naturalization of alien succulent plants in New Zealand, but also accounts for my
personal evolution on the matter. I have also omitted to mention this plant in my 2007 Romanian article Plante suculente narturalizate in Noua
Zeelanda published on Aztekiums website thinking in error that it has to be a native succulent.

(2) Succulent plants from down under Sarcocornia quinqueflora an odd succulent halophyte (2007).

(3) Referred usually by botanic authors as an annual plant, reportedly in some situations it may have a biannual or even short-lived perennial habit,
with some shoots remaining leafy during winter. Such bizarre alterations of the habit have been seen in another halophyte plant naturalized in New
Zealand Salicornia europea. John J. Knapp, Joseph M. DiTomaso & al. (2003) are the only botanical authors I have read who consider the plant as
being biannual, Flora of New Zealand Vol. 4 mentions only the leafy shoots remaining during winter.

(4) I wont enter into taxonomic details; I will only mention that the well-known Cakile maritima ssp. integrifolia often mentioned in British botanical
records is actually a synonym of Cakile maritima ssp. maritima.

(5) The late Dr. Phil Maxwell (the author of The Rhipsalis Riddle - or the day the cacti came down from the trees, 1998 I highly recommend this
article) was a very fierce critic of the easy-explanation long range dispersal vectors (birds, sea currents, land-bridges connecting everything between
actual continents to mythic land-masses, etc.) usually advocated when no obvious reason is available and I totally agree with him. However, it was
proved that Cakile maritima is able to be dispersed on very short distances by the joint action of tides and coastal currents and therefore I cannot rule
out completely in this case this dispersal vector even on longer distances.

(6) You cant miss it when in flower the flat top of the racemes is characteristic for the species.

(7) It was due mainly to the leaf form and design that we have many names and synonyms given to his plant, mainly in the 18
and 19
There are four different factors though a natural variability of the leaf form, spots arrangement and colour, flower colour and (very visible in
cultivated plants) the intensity and duration of the sun exposure allowing for different looks.

(8) I hope that I will be able to write some time next year about the Opuntia plants of Australia, their invasive behaviour and the surprising lack of
success in New Zealand.

(9) On Didge Rowes Xerophytes NZ notice board David Kirk of Tauranga wrote: As a kid I remember a monacantha 'tree' growing next to a right-of-
way at Mt. Maunganui, a massive spiky trunk and all... I wonder if it's still there? Walking in the sand dunes was sometimes relatively perilous, as the
dunes were a favorite dumping-ground for garden waste and small bushes of the Opuntia were established in the undergrowth. They've all gone now,
but there remains an interesting mixture of old-fashioned bulbs I've never seen elsewhere (apart from two Lachenalia sp); at least three unknown
Gladiolus species, etc., and the commonly-encountered Cotyledon orbiculata v. orbiculata (?), anyway, the silver-grey one.

(10) Well established on sand dunes and growing amongst Opuntia ficus-indica, Lupinus arboreus and Ammophila arenaria.

(11) Growing on rock rubble with Opuntia robusta, Sedum dasyphyllum and Ornithogalum longebracteatum.
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All errors, omissions and misconceptions are mine.

All photos by Eduart Zimer, except 6 by Dan Georgescu.

Eduart Zimer, September - October 2008
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