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Succulent plants from down under Adventive Plants (Part 8):

The bid for succulence: Calystegia soldanella - Bits and Pieces (Natures potting mix for succulents /
New pictures)
XIX. The bid for succulence: Calystegia soldanella
Before we start I have to clarify few things the well known Calystegia soldanella (Linnaeus) R. Brown 1810
(basyionym Convolvulus soldanella Linnaeus 1753, heterotypic synonym Calystegia sepium var. tangerina
Pau 1924), which is in fact an invalid name (1), correctly being referred as Calystegia soldanella Roemer &
Schultes 1819, is actually not a naturalized or adventive, but a cosmopolite plant spread in both hemispheres. It
belongs to Convolvulaceae, the Bindweed family. Calystegia soldanella is indigenous in New Zealand (North
Island, South Island, Kermadec Islands, Three Kings Island, Stewart Island and Chatham Islands). Beside its
English widely used common names like Shore Bindweed, Sea Bindweed, Sand Bindweed (or Convolvulus in
some variants) or beach Morning Glory this plant is also known here by its Maori names of Rauparaha, Pohue
and Nihinihi. Therefore Calystegia soldanella is not a naturalized plant, but is not exactly a native either (2) (3).
Basically it grows in both hemispheres mostly in temperate to temperate with sub-tropical influenced oceanic
climates. To make it worse for my decision to include this plant into an account dealing with New Zealand
naturalized succulents it is not even a typical succulent. In other words if you are strictly a succulent freak,
please skip this chapter!

1. A splendid young
soldanella growing in
pure sand in Opoutere
(Coromandel Peninsula).

Calystegia soldanella is a glabrous perennial herb with creeping rhizomes and slender much branched aerial
succulent stems, mostly trailing, rarely twining, and forming usually small patches which can reach in some
cases over 200 cm across. The petioles are usually 10 60 mm long, occasionally up to 80 mm long, and
slender, in most cases longer than the leaf. The leaves are (10) 15 50 (80) (10) 20 50 (75) mm, usually
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sub-succulent, in certain circumstances becoming either membranous or succulent, glossy, covered with a thick
waxy protective coating giving the leaves sometimes a leathery aspect, entire, more or less reniform (4) with a
usually shallow and rounded sinus, with the base often broad cuneate; the apex varies quite a lot from clone to
clone from emarginated to rounded to acute. The flowers are solitary and pleaded and folded in bud. The
ribbed peduncles are usually 15 70 mm, on occasion up to 100 mm long (longer than the petioles), the bracts
are broad ovate, obtuse, (10) 12 15 (18) mm long, the sepals are obtuse, very similar to the bracts. The
corolla is pink to purplish, mostly with white mid-stripes, occasionally all white, 20 40 x 25 50 mm, funnelshaped (campanulate). The ovaries are surrounded at the base by a nectar-secreting disc. The capsule is ovoid
to broad ovoid, 1215 mm long, apiculate. The seeds are dark brown and smooth. The flowering period is
October to The flowers are hermaphrodite (have both male and female organs) and are pollinated by bees,
moths and butterflies. March and is fruiting in March in the southern hemisphere of course.

2. Calystegia soldanella
in Wreck Bay, in the
Rangitoto Island. The
substrate is a bit
different; the plants seem
to prefer the limit of the
beach where the lava
fields begin. Note the
accumulations and the
moist sand.

Calystegia soldanella can be easily distinguished from other similar species due to its kidney-shaped leaves,
but it can form also hybrids with indigenous and introduced Calystegia species (namely Calystegia
tuguriorum), the hybrids being also easily distinguished by the sub-succulent stems and leaves subsucculence being not an ordinary feature of the Convolvulaceae family and by the intermediate leaf form,
oscillating between reniform and deltoid (as in the majority of the Convolvulaceae). It is also suspected that
natural hybrids with Calystegia sepium ssp. roseata and Calystegia marginata may occur in New Zealand,
although this hasnt been documented yet (New Zealand Plant Conservation Network).
Being a cosmopolite species no wonder that it is also polymorphic. The polymorphic character relates not only
to general morphologic features having a zonal representation, but very interesting particular habitat
conditions can generate different growth forms. However, most of the morphologic variations based on genetic
differentiation analyzed by Takanori Ohsako & Gakuto Matsuoka (2007) in Japanese shoreline populations
isolated from inland populations for up to 800,000 years (5) have occurred not between individuals from
different populations, but 80% of them occur within the same population.
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3. Calystegia soldanella
flower detail of one of
the Opoutere plants.
These plants are not very
prolific flowerers, but
note the number of buds
present in a small area.

4. Larger patches of
Calystegia soldanella
growing in pure sand
on the beaches of

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The most interesting fact is that Calystegia soldanella usually growing on beaches or fore dunes in
xerophytic conditions has very fleshy, succulent leaves, capable to store a certain amount of water in order to
overcome the draught; upslope plants growing more or less hidden between higher vegetation in moist and
sheltered positions have membranous leaves (more like the other Calystegia or Convolvulus species have). It
also can re-convert in a typical climber, usually on scrubs, in search for better light. In other words this plant
has definitely placed a bid for succulence, but seemed to have stopped halfway through, becoming a static,
slow to non-evolutionary species at least seeing habitat pictures of Calystegia soldanella from all over the
world this is my gut feeling. The interesting thing is that it can easily revert (so to speak) to membranous
growth forms depending on the habitat characteristics; it seems to have kept a copy of previously functional
genetic setups. I havent seen plants with membranous leaves but it was obvious to me that plants growing in
more hostile conditions have also fleshier leaves. On top of that its thick, fleshy roots are also designed to store
precious water and nutrients. More, the waxy coating of the leaves prevent excessive perspiration. Definitely,
we have enough features to which could be regarded as a bid for succulence, even if marginal in this instance.

5. Leaf detail of a
plant from Wreck
Bay in Rangitoto.
Note the typical leaf
glossy appearance
and the tiny apical
soft spike you dont
see in all plants.

But despite all these remarkable adaptations Calystegia soldanella seems to have stopped its evolution toward
succulence, abandoning the evolutionary traits followed independently by other plants. The most relevant thing
is that Calystegia soldanella still uses the usual C3 photosynthesis (6) most plants have, and this is definitely
not that common in succulent and xerophytic plants. For a plant with xerophytic adaptations C3 photosynthesis
brings a lot of disadvantages like photosynthesis taking place on the entire leaf surface and only on the leaf
surface and only during the day (therefore all conditions for a high perspiration rate are present). C3
photosynthesis is more efficient under cool, moist and moderate light conditions and also, by being the first
mark of this remarkable physiologic design which is photosynthesis, has maintained the basic but very
functional physiology involving less engineering to make all this work fewer enzymes and no specialized
anatomy (7).
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6. Trailing stems on a
rocky substrate in
Wreck Bay (Rangitoto
Island).Compare the
leaf form of this plant
with that of the plant
in picture no. 5, from
the same location, just
20 or 30 meters away.

soldanella growing
in pure sand in
Raglan (Waikato),
at the Tasman Sea.

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Although it has outspoken xerophytic adaptations Calystegia soldanella always prefers the vicinity of the
water; as it has a moderate salt resistance it usually grows in coastal habitats, but also sometimes inland along
lake shorelines (8). It prefers moderate dry to slightly damp sandy or shelly beaches, banks, sand dunes or fore
dunes, but is relatively common also in drier gravely or rocky areas (as I have seen in Rangitoto), advancing
even occasionally in coastal turfs or on cliff faces, or even up to 30 meters upslope in grassland. Especially in
sand or very open sandy soils the stems run under the surface with leaves emerging from the sand; what seem
to be different plants could be one and the same.
However, it requires a certain amount of moisture in the air or substrate deep down under the surface of the
sandy beaches there is also moisture available and definitely good light (it simply cannot grow in shaded
positions). In its typical state Calystegia soldanella is a sand dune pioneer, establishing quickly in exposed
positions and becoming locally dominant, but less inclined to face the competition of other plants. It is
regarded as a very useful sand-binding plant due to its rhizomatous habit and matted, trailing stems (Flora of
Tasmania online).
Like many of the bindweeds, Calystegia soldanella is also known from cultivation. It makes a wonderful
display when placed in full sun, in sandy or rocky well draining soil. It accepts anything from slightly acid to
neutral to slightly alkaline soils, but does not like too much organic compounds. It is very easy to propagate by
fresh seed sown directly in situ in cooler periods of the year (and if you have enough patience as germination
can take 1 3 months at 15 degrees Celsius) or in a cold frame if freezing winters in your area, cuttings made
in summer (lay them simply on the ground and cover the cut area with sand) or division of the rhizomes very
efficient when made in early spring while the plant is still dormant and especially when seed grown plants are
divided. I imagine it would make a nice display with its long lasting showy flowers and green glossy leaves. It
is also quite hardy; adult plants can stand moderate frosts, but not seedlings. But even if it looks to be one of
the toughest, hard to kill plants, actually it is not. It usually is quite difficult to establish in typical gardens
because it simply does not like heavy and rich substrates, but given the right conditions you cant stop it! Once
established, you have just slim chances to eradicate the plant, as it starts again from seed, rhizomes or rooted
stem fragments.
Calystegia soldanella is not only (potentially at least) a nice garden plant, but has also in a certain extent
medicinal uses. It has a well known purgative effect (oh well, say side effect), in fact the Plants for a Future
database lists antiscorbutic, diuretic, febrifuge, irritant, purgative and vermifuge properties.
The plant has also culinary uses. Young shoots are cooked as a vegetable or pickled and used as a samphire
substitute I imagine it cant be a gourmet meal even if we speak here of international cuisine, but still it used
to be an ingredient in survival meals. The New Zealand Plants Conservation Network gives also a quite
peculiar use: The Maori gathered the thick, white, fleshy roots and pounded these to form a pulp; this was
then used as a relish to flavour some meats.
The economical importance even if marginal does not stop here; again Plants for a Future database: The
stems are very flexible and are used as a string for tying. Fairly strong but not long-lasting.
I have seen Calystegia soldanella in several locations (it is common, not threatened), but always on the
seashores, growing in anything between pure sand to stabilized sand dunes, except the Rangitoto plants,
growing on rough gravel in the northern parts of the island (namely in Wreck Bay). It is a plant you usually
dont notice, its not striking if not in flower. But at some stage in December 2007 during a visit at the
Opoutere beach (Coromandel Peninsula), a splendid and little altered coastal habitat, I have noticed the
uniqueness of this plant growing in pure sand. It is not present in large numbers, but you can see here and there
patches of Calystegia soldanella with leaves rather on the succulent side, as it is a quite dry environment here,
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especially in high summer. The other well represented plants here are Cakile maritima ssp. maritima (again in
its succulent version) and Spiniflex sericeus. In a similar habitat at Raglan (Waikato) Calystegia soldanella
grows in pure sand as well, but being on the west coast at the Tasman Sea I reckon it gets more rain than the
Coromandel Peninsula plants.
At Piha (west of Auckland) Calystegia soldanella grows on very exposed beaches westerly winds spread fine
particles of salty sea water well away from the waterline but also on the fore dunes again accompanied
usually by Spiniflex sericeus.

8 9. Another flowering plant in Opoutere

(below) and a very nice relatively modern
drawing of a Calystegia soldanella extracted
from a 20th century herbal written by M. Grieve
A Modern Herbal (1931) (right).

The Rangitoto plants grow on a different substrate. As there are no sandy beaches in Rangitoto Calystegia
soldanella had the only choice to accommodate on the rocky or gravely shores of the island. As the seeds are
quite salt resistant the dispersal occurs by sea currents, waves and tides. I imagine that being close to the
mainland Rangitoto would be a target hard to miss. It was very interesting to notice the differences in leaf form
displayed by plants growing very close to each other. Unfortunately, my targets for the trip were set and days
are never too long on Rangitoto, I took just few pictures and continued my way.
I have to admit that I wasnt interested in this plant until the austral summer of 2007/2008, when I saw the
flowering plants on the Opoutere beach. But as a matter of fact, even if most of the botanists and plant
enthusiasts do not regard Calystegia soldanella as a succulent plant, I think this plant has definitely placed a
bid for succulence at some stage in its early evolution. Evolutionary convergences with known succulent
species are easy to observe a certain degree of succulence (sub-succulence in botanic English, Id like to call
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it marginal succulence) of the stems and (especially relevant) of the leaves and the waxy coating of the leaves
which prevents perspiration and protects the plant against lashing winds and sea water mist.
XX. Bits and Pieces
Natures potting mix for succulents
We worry much too often about how much grit or sand or humus or moisture retainers or whatever compounds
shall we add to the potting mix we prepare for our cacti and other succulents. Of course, you cant compare
potted conditions with free root run conditions in gardens and none of these with the challenges of life in the
wild. The amazing thing is that only looking around at wild living plants you can actually understand how
much they can take, and how little they need to be pleased. I find it extremely useful to take some time and
look at the substrates some succulent and non-succulent plants grow in nature on. The following pictures (some
of them being already published elsewhere) are only a very small glimpse of natures secret potting mix recipe
for succulents. Im not prepared for a lecture on this, but just have a look and have some thoughts.

10. Some succulent plants are able to thrive in (or only in) pure sand like this Cakile maritima ssp. maritima from Opoutere
(Coromandel Peninsula). The coastal sand dunes and beaches environment is actually not that hostile as you think it would be and
even if the surface becomes extremely hot in high summer just few centimeters below the surface the deep roots stay in a relatively
cooled substrate. Sand also has good water retention and 30 or 40 centimeters deep there is always some moisture available.
There is a downside though a certain degree of salinity associated with sometimes high Calcium content (given by the crushed
shells). And by the way - in most of the cases the total lack of organic nutrients makes them fight hard every single day.

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11. In estuaries or salt

marshes margins towards the
sea, where the sand is usually
wetted periodically by tides
or waves with a certain
dynamic, pure sand is often
replaced by a mixture of
sand, shingle, rocks and
crushed shells like this
succulent chenopod) from
Miranda grows on . When for
some reasons the area is
permanently exposed more
plants can establish. A good
candidate for a succession
plant is Carpobrotus edulis.
Of course, the easy-to-get
problem hasnt
been solved yet. Once left to
dry out this kind of substrate
improves its drainage.

12. This substrate Sedum

album from Rangitoto is
growing on looks similar to the
one above, but actually it is not.
The main difference is given by
the rocky base (lava fields in
this case) and a certain
elevation above the sea level. It
is definitely much drier, but it
contains more or less the same
mixture of sand, shingle, rocks
and shells. Whatever the reason
is high seas, storms, flooding

sediments is usually reasonable
superficial and dries out fairly
quickly even after torrential
rains as water has nowhere to
accumulate (except for small
pockets) and simply flows
depending on the particular
conditions and more or less
decayed organic debris can
form a layer or mix with the

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orbiculata var. orbiculata
growing on Lion Rock, Piha,
west of Auckland at the Tasman
Sea. This is a high rocky
colossus in the middle of the
beach and in some places a
considerable accumulation of
sand and clay particles brought
by the strong winds has been
formed on milder slopes.
Humid air from the sea has
helped this soil to compact and
later on emerging vegetation
has added organic debris and
decayed matter, roots have
bound everything together. A
thin layer of soil has been
formed over a long period of
time, being the life support for
varied vegetation forms. I am
not a geologist but it is clearly
that this soil has little in
common with the surrounding
beach and fore dune areas.

14. Carpobrotus edulis the

yellow flowering form at
Tahuna Torea (Auckland).
The substrate is similar to
that in picture no. 11 but the
site is just a bit more
elevated. Dense mats of
vegetation allow organic
debris to accumulate and
decay, eventually changing
the texture of the soil. When
the conditions come right,
succession vegetation will
take over forcing pioneers
like Carpobrotus edulis to
seabed has been for some
reason uplifted and became
permanently exposed. Of
course, there are a lot of
factors which can derail this
process and change the
characteristics of the site, but
if left alone and this can
also happen in only few
decades we may find on the
same spot pampas grass

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15. This fine Kindergarten for

succulent plants tells us a
different story. The substrate is
different from the picture no. 12
because it has less fine sand
particles more traces of
volcanic ashes (which is not
necessarily very fine), and a
thin crust of pebbles and
gravel. This type of substrate
seems to work very well for
young Aloe maculata and
robusta. Under and between
these medium sized particles
available for some time, where
seed, leaves or rhizomes have
just enough to create new
plants. This kind of substrate
does not occur on large
surfaces but is rather restricted
to smallish patches between
lava blocks.

16. Compared to the previous

picture this is a step
Rangitoto use very wisely the
fissures or the small crevices
between lava blocks, where
accumulations of volcanic
ashes or smaller rock
moisture. In this particular
Agave colony there are no
higher vascular plants, but
lichens, mosses (in moist
positions) and xerophytic
grasses are growing instead
from the smaller sized
pockets of soil.
In this
particular case the above
mentioned vegetation types
create a more composite
vegetation pattern of the
ground layer.

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17. Well, I should have started

probably with this picture. The
typical vegetation succession is
bare rocks lichens small
vascular plants, but in shaded
places mosses can cover the
ground almost completely. The
substrate in this picture is
similar to that in picture no. 15
but the vegetation development
path will be very likely a
different one. Mosses retain
moisture and in time dead and
decayed plants will create a
thin layer of fertile organic soil.
Instead of succulent plants
there will be probably fast
growing non-wooden high mass
vegetation established here in
some time, changing and
preparing the environment for
a repeated succession. As this
picture was taken close to the
bush limits it will be eventually
conquered sooner or later by

18. This eroded coastal

interesting because it clearly
shows how thin the fertile
layer of the soil sometimes
can be. On a thick layer of
sandstone a bush like
vegetation has established for
a long time, it could be many
hundreds of years. However,
probably nowhere in this
area the fertile soil layer is
deeper than 20 40 cm. It is
most likely the norm here on
elevated platforms. Although
the only succulent plant I
could see here was Crassula
multicava, I think there are
virtually good chances for
other succulents to establish
on the borders of the dense
vegetation where the soil is
relatively often disturbed.

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19. This rocky outcrop in

Gulf), how hostile it may
look, is still home for several
plants Disphyma australe
ssp. quinqueflora in this
platforms have always some
sort of cracks, or fissures, or
moisture can be available for
the plants. Nick Perrin (the
editor of the CSSNZ Journal)
and the author of an 1996
australe ssp. australe) told
me once that these small
endemic mesembs can send
their roots up to 2 meters
deep following the cracks of
the rocks in search of
precious moisture.

20. This is the image of a

wet sandstone rock on
Rakino Island (Hauraki
sedimentary rock its
texture is prone to form
cracks and fissures when
exposed to the natural
elements. It is also very
porous and keeps traces
of moisture even deep
inside. Sandstone is a
preferred substrate for
many xerophytes and
succulents because it
completely deep inside;
it is for the plants just a
matter of sending their
roots and stay connected
to a minimal source of
moisture which can be
the difference between
life and death in times of

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21. Vertical cliff faces are

not a problem for some
plants like Cotyledon
orbiculata var. orbiculata
growing on a tiny lodge on
Lion Rock (Piha). The big
advantage is that even in
humid climates because the
rainwater flows away
instantly xerophytes can
have their portion of
draught. This particular
spot knows also excellent
natural ventilation; strong
are battering
almost continuously the
area, which is again for
Cotyledon orbiculata var.
locations here in New
Zealand I imagine this is
the optimal habitat for this
type in South Africa as

22. It is not always a matter

of cliffs and sandstone and
fissures and cracks, or of
retaining moisture for the
days of draught to come that
succulent flora. This rough
lava field few hundred meters
from the southern wharf of
battered by winds laden with
sea water. When this
particular condition occurs,
you cant expect too much
but only halophytes to
establish like Agropyron
junceiforme grass in this
particular case. I havent
seen Disphyma australe ssp.
australe here (another good
candidate) but patches of
present close to or within the
splash zone.

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23. A sandstone cliff in Russell Bay of Islands (Northland) in a spot with strong marine exposure bearing some
vegetation (the small tussock in the centre is Sarcocornia quinqueflora ssp. quinqueflora). In more sheltered
positions an almost continuous strip of Crassula multicava is growing at 1 meter above the high tide mark.

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24. Well, I have captured

prerequisites we need to
make my point we have
the porous sandstone rock,
we have the fissures and
pockets, and we have a
very healthy Disphyma
australe ssp. australe
making the best out of a
given situation on Rakino
Island (Hauraki Gulf). It is
not a singular occurrence,
but several plants were
scattered on the rock
surface, some of them of
flowering size and bearing
numerous seed pods. This
is a detail of the rocky
outcrop seen in picture no.
17. As you can see lichens
have tried as well to make
a living here.

25. A similar example is

given by this picture this
time at a certain distance
from the sea in Rangitoto a
tiny seedling tries to establish
on rough and fragmented
lava fields and eventually it
will succeed. There is a
difference though at this
distance (possibly 150 200
meters) from the sea and
somewhat sheltered by some
bush patches placed in
between, there is no cooling
breeze, but an incredible heat
emanated by these basaltic
lava fields especially in high
summer during sunny days.
Reportedly the rock surface
reaches 70 degrees Celsius,
hot enough to cook some
eggs. The survival of this
improbable, but hey - this
picture was taken in high
summer! Also remarkable is
that here plant colonization
has just begun.

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26. In similar conditions

tetragona ssp. robusta on
the lava fields of Rangitoto.
The difference is that this
particular spot lichens are
already very established
and some other plants have
also tried their luck I the
past. Again, the plant is
speculating perfectly the
opportunity given by the
moisture retained in the
cracks and the possible
traces of organic matter
generated by the decayed
vascular plants. Without a
proper organic layer the
heat is almost unbearable
in this open space not very
close to the sea.

27. The ultimate success

a small plantlet has
established in a narrow
crack in the rock,
surviving killer heat and
heavy draught for years
and years, growing
eventually into a big
tree! It is kind of a
miracle, as there is no
groundwater here in
Rangitoto. It also proves
that no matter how
harsh the environment,
no matter how big the
losses there are always
winners in the vegetal

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New pictures
As a bonus for this issue rich in illustrations here is a selection of 9 pictures of naturalized succulent plants I
havent published before.

28. A small Agave

americana in Rangitoto
from the small colony
halfway between the
Islington Bay wharf and
the passage to Motutapu
Island. By the time I took
this picture (January
2009) the colony was
checked and all bigger
plants removed or (in
two cases) simply half
uprooted and butchered
on the spot. However,
several smaller plants
from 30 - 40 cm plants to
small plantlets like this
one still exist.

29. This is the flower of

fluminensis - also called
The Wandering Jew for
the rapid spread it can
achieve a creeping
groundcover with fleshy
leaves and succulent
stems. I took the picture
southern end of the
beach, where the vertical
cliffs start. This is a hot
plants can be seen.

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30. Thats me with an

unidentified Agave sp. in
Mellons Bay (Auckland). It
is a survivor plant, not
because of the harsh
conditions, but because of
the sheltered, moist and
position, surrounded by
dense vegetation which is
even worse! Probably it
originates from garden
waste disposed on the
beach; I couldnt see any
other Agave on the beach
or on the higher grounds in
the back. To be honest, I
didnt put too much effort
in identifying this plant;
however, I am convinced
that it definitely has an
untypical growth.

31 - 32. Details of the unidentified Agave sp. in Mellons Bay.

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33. This Aloe arborescens is not exactly a wild plant, but still survives in an abandoned
garden in Western Springs. I have inserted this picture more to allow a direct comparison
of this neglected plant with the true survivor specimens of Rangitoto (see in Part 7).

34. Carpobrotus edulis at

Tahuna Torea (Auckland),
a small salt marsh I have
already mentioned several
times before. Succulent
plants have a limited
occurrence here (the other
succulent plant I have
quinqueflora) but still it is
a wonderful piece of

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Gazania rigens on
the Port Waikato
fore dunes. These
have been massively
eroded in the past,
so that basically any
type of vegetation is
welcomed. It is not
quite conservation
but it is a start.
amazing thing is
that nature took
course on its own.
At this stage the
closed for camping
and provides only
access with the
results you can see.

36. Yucca gloriosa

marginal succulence
growing on the
fore dunes of Port
Waikato. There will
be a short account
on this plant in the
months to come
take this as a

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Additional References:
African flowering plants database;
An Encyclopedia of New Zealand, 1966;
H. H. Allen Flora of New Zealand (Vol. 1, Government Printer, Wellington, 1961);
H. H. Allen Flora of New Zealand (The updated electronic version, Vol. 1, 2004 - );
Flora of Tasmania online -;
M. Grieve A Modern Herbal (1931);
The International Plant Name Index -;
Native Plants at Piha -;
Takanori Ohsako & Gakuto Matsuoka - Nucleotide sequence variability of the ADH gene of the coastal plant Calystegia soldanella (Convolvulaceae) in Japan
Kathleen Robson, Alice Richter, Marianne Filbert - Encyclopedia of Northwest Native Plants for Gardens and Landscapes (2007).

Further Readings:
Ute Drobner & al. The sand dune vegetation of Chrystalls Beach, Southern New Zealand, with particular reference to the cushion community;
Frances C. Duguid - Botany of northern Horowhenua lowlands, North Island, New Zealand (1990);
A. E. Esler - Botanical features of Tiritiri Island, Hauraki Gulf, New Zealand (1977);
Plants for a Future -;
C. J. Webb, W. R. Sykes, P. J. Garnock-Jones - Flora of New Zealand (The updated electronic version, Vol. 4, 2004 - );

My Notes:
(1) A comment from The International Plant Name Index: Brown did not make this combination on p. 483; his citation of "C. Soldanellae L." on p. 484, if
considered as Calystegia, was still only provisional. In a discussion clearly concerning Calystegia reniformis Robert Brown compares it with C. soldanella L.
wondering if there were distinct species: "Obs. Facies C. Soldanellae L. diversa praecipue pedunculis teretiusculis: an vere distincta species?" Even if we could
interpret that C. soldanella means Calystegia soldanella and not Convolvulus soldanella but indicating Linnaeus as authority this assumption becomes
extremely doubtful the combination Calystegia soldanella (L.) R. Brown would be invalid and has to be regarded as a personal comment simply because
Robert Brown did not expressly indicated that he accepts or promotes a new combination. However, some authors consider the two names being synonyms. The
generally accepted nomenclature is:
Calystegia soldanella (Linnaeus) Roemer & Schultes 1819.
Calystegia soldanella (Linnaeus) R. Brown 1810, nom. inval.
Calystegia soldanella (Linnaeus) Choisy 1845 (?).
Convolvulus soldanella Linnaeus 1753 (basionym).
Calystegia soldanella var. australis Endlicher 1883, nom. illeg.
Convolvulus sepium var. soldanella (Linnaeus) C. Moore & Betche 1893.
Convolvulus sepium var. saldonella F. Mueller 1875, nom. inval., nom. nud. Note saldonella (sic).
Calystegia reniformis R. Brown 1810.
Convolvulus reniformis (R. Brown) Sprengel 1824.

(2) Even NZ authors have a dual position i.e. by being referred in both vol. 1 (indigenous plants) and vol. 4 (naturalized plants) of Flora of New Zealand.
(3) We cannot be even sure that native stock has not been mixed in the past with overseas plants, as it happened to other relatives. See Calystegia sepium
(Linnaeus) R. Brown ssp. roseata Brummitt 1967 x Calystegia silvatica (Kitaibel) Grisebach ssp. disjuncta Brummitt 1967 ex
(4) Reniform means kidney shaped.
(5) Since Calystegia soldanella does not have long-distance terrestrial dispersal systems (Takanori Ohsako & Gakuto Matsuoka, 2007) inter populations gene
exchange is not a credible explanation for the relative uniformity. However, I think that based on the global distribution of the species indicating an ancient

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origin the isolation period of the populations analyzed by Takanori Ohsako & Gakuto Matsuoka considered to be mostly between 500,000 800,000 years is
not long enough to be responsible for a relevant genetic differentiation in a such static, old timer species.
(6) It is called C3 because the CO2 is first incorporated into a 3-carbon compound.
(7) For example there is a single enzyme (Rubisco) involved in both photosynthetic processes and CO2 intake.
(8) For New Zealand Lake Taupo is a very good example.

-----------------------------------------------All errors, omissions and misconceptions are mine.

Eduart Zimer, August 2009

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