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Succulent plants from down under Adventive Plants (Part 3):



What drives the success of the aliens? Carpobrotus edulis, a friendly alien?



VII. What drives the success of the aliens?

It is hard to define the success of alien plants if we just say that it is the ability to occupy a certain
habitat and to coexist with, limit the distribution of, or simply to eliminate a competitor plant this
would mean that we have grossly oversimplified things. A. E. Esler (1988): Considerable success is
accredited to those which regularly coexist with other species, or occupy temporary or difficult
habitats. In fact to measure and appreciate the degree of success is a very difficult task as success
can be only apparent or temporary and there are enough examples in succulent and non-succulent
flora of New Zealand. Alien plants may escape in nature and form strong populations in a short time
but that is just the beginning as the harder task is to maintain and expand those populations; thus for
some types of plants the relative number of young plants and seedlings in a population is an
indicative of their success at least for the time being (or better said of their potential). This does not
necessarily grant their success as even some of the most annoying and persistent aliens (as Cirsium
vulgare, even if not a succulent plant) have a high mortality rate of young seedlings. On the other
hand we usually have a static view of this phenomenon and records made at some point in time
might be totally irrelevant just few decades later. Time is as always a critical factor. There are quite a
few plants mentioned in early botanical records in New Zealand which couldnt really make it over
time, some are known just from casual collections many decades back and havent been seen since or
simply remained very limited in distribution or number of individuals. Opportunities given and
available habitats have changed in time - A. E. Esler (1988): The observations of Kirk (1871) and
Cheeseman (1883) serve as a base line for judging the performance of alien plants present in
Auckland from that period. It has to be appreciated that opportunity has changed meanwhile. The
small urban area had unpaved roads and rough footpaths; the rural areas were extensively ();
pasture management was less able to cope with weeds; there were no selective herbicides. Some
alien plants that thrived then have little opportunity now; and the reverse is also true.

Not of a lesser importance is the fact that although for most of the vascular plants the vegetative
reproduction is a secondary form of dispersal, for the succulent plants this reproduction form is
usually very effective and works very well especially in new habitat types, where specific pollinators
do not exist, or the needed germination and / or growth conditions of the seedlings are not met
regularly or in all years. This reproduction type is generally the only available for hybrid plants. This
is a very important feature as some of the plants escaped from cultivation, even if true species and
not hybrids, might have been previously cloned in cultivation and are not capable of sexual
reproduction. In this case the vegetative reproduction is their only option. On the other hand there is
a flop side too vegetative reproduction does not generally favor a wide dispersal but just a localized
or even a very restricted distribution. There are of course several noticeable exceptions in New
Zealand succulent adventive plants and probably the most important of them is the well-known
Crassula multicava (1).

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The behaviour of new introduced plants is not entirely predictable and in most of the cases even if
there are known difficulties experienced in other countries it is still hard to assess their future
development in new habitats. This takes time (meaning many years or even decades) therefore it is
even harder to assess correctly recently arrived plants. Every species has its unique way to success
during which several encouraging or discouraging factors are in fragile balance with its natural
abilities and opportunities given. Here are some of the main discouraging factors (2):

- The adaptation effort. The natural habitat in which the alien plants end up is not necessary
similar to the original native habitat, beyond all appearances. A desert for example is not any
desert its not like Atacama, Sahara, Kalahari, Mohave or whatever; in fact there are totally
different habitats beyond evident similarities. A new habitat may have some of the
characteristics of the original habitat allowing in a first instance and under ideal
circumstances alien plants to establish and to disperse but still there is an adaptation effort
necessary, and not all newcomers have on a long run the ability to adapt to the new
conditions. We have to understand that there are also very important invisible factors as the
specifics of a biologic system starting with the most simple and primitive organisms and
ending with the vascular flora or with the characteristic fauna. It is not necessary easy for an
alien plant to adapt to this new biotic frame; most of them do not succeed in this attempt (3).
The particular climate may also have different patterns and unexpected twists with hard to
foresee consequences, which may occur from time to time. A certain plant, incapable to
penetrate in a compact and hostile habitat, may have a window of opportunity - say a massive
landslide, which is an open invitation. It may flourish for some time without any competition
of any kind; it may establish self-sustainable populations in a short time; it may benefit few
years from an adequate climate. But for most of the challengers this adventure will end very
likely in few years, after a freezing winter, or after several wet years, or hot years or dry
years or decimated by competitor plants with greater dispersal capabilities building up their
populations, or by diseases (generally viral or fungal) for which the plant has not developed a
natural immunity or long term survival means, and so on. It is not always nice and easy to
be the new kid in town.

- Restricted initial dispersal and initial habitat type. The very most of the escapes (at least
in recent years) are of horticultural origin, usually from garden waste, abandoned gardens but
also seed or vegetative dispersal beyond the intended areas. The initial dispersal takes place
usually within or very close to urban areas; these are temporary and ever-changing habitats,
which at some point may know a drastic change or may even disappear. As city habitats are
becoming in New Zealand less and less friendly for uncontrolled vegetation spread this
means that there is less and less time available for the alien plants to make it in the outer
natural world. This may explain very well why several of the plants collected from urban
Auckland area in the 19
th
century are nowadays rarities or have simply disappeared.

- Vegetative reproduction. I have already mentioned this; usually it allows their survival but
stands just for a very restricted distribution. The succulent plants naturalized in New Zealand
are lacking one of the dispersal mean other plants are using very efficient easily detaching
segments or stems getting caught in the fur of wild animals as the lack of this kind of animals
as well.

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- The lack of specialized pollinators. Some plants are relying on them and cannot be
pollinated by other means. Yucca Tegeticula maculata (or Tegeticula yuccasella) is a very
well known example. As usually imported plants come not in family packs with their
favourite pollinator this is a serious handicap. It was also stated in botanical literature that in
some plants not depending on specialized pollinators the flowers were not recognized by
the native insects and therefore widely ignored, in the end with the same effect. In this case
an almost pure theoretical chance exists wind pollination, but the succulent plants we are
speaking about have extremely slim chances because they do not produce pollen in high
quantities and usually individuals are too far from each other.

- A sustained vegetation control. Environmental and ecological organizations are becoming
more and more effective in controlling vegetation growth and dispersal of alien plants,
animals or insects, especially in reserves, iconic habitats and natural landmarks.
Unfortunately the damage was done long before a pro-active habitat protection was
acknowledged as a high priority.

- A much reduced genetic pool. Usually the cultivated and naturalized plants are not even
close to the genetic variety existing in the original habitats. Some of the escapes are
originating from cloned material or were already hybridized or are the result of artificial
selection. On the long run this is a serious handicap.

- Last but not least the limited compatibility of the biotic systems. In some extent the
gondwanic nucleus of New Zealand is favouring more plants originating from ex-gondwanic
land masses (such as Africa, South America, Australia, India and Madagascar) and less plants
originating from the northern hemisphere. The composite origin of New Zealand (a
gondwanic nucleus surrounded by foreign terranes, resulted from lifted ocean floors) is
however a limiting factor by itself, this explains why even Australian plants find sometimes
very hard to set foot here. To be fair most of the succulent invasive plants are originating
from South Africa. This limited compatibility is far from being a rule of thumb but observing
and analyzing the distribution of several plants throughout the world might reveal its
significance.

Its not all about handicaps but also about the natural abilities of the plants in ideal conditions, the
behaviour it is usually displayed and which can make the difference. Their behaviour can be
classified as follows:

a) Plateau species. After the initial dispersal and growth of the populations, the number of
individuals or their density does not exceed a certain level. This level can be low - usually
just few individuals or small populations scattered over a wider territory; medium - the
growth period is longer and the rate is higher, they also manage to establish and maintain
significant populations with a reasonable high density and in a certain extent to integrate in
the new habitats; even if quite frequent the individuals are not in such great numbers to alter
the habitats or to endanger or to exclude competitor plants. This is the highest level the very
most of the alien plants have ever reached. There are few high plateau species though which
can cover wide areas with high-density populations. Although a certain level cannot be
surpassed this is in its extreme case the most dangerous level as competitor plants can be
easily replaced and habitats changed in a manner which may produce further changes in the
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structure and dispersal of the native flora and fauna, triggering in worst case unforeseeable,
uncontrollable and irreversible alterations.


b) Increasing species. The main difference is that the plants are not building rapidly towards
their maximum potential expansion, but in a somehow slower motion. However, it shows the
ability of the plant to overcome environmental pressure working actively against them. The
increasing rate can be slow, medium or fast. The slower the growth rate of the populations is,
the better the chances are to fight or eradicate invasive plants. Fast increasing species are
always a bit of a worry because there might be not enough time to act consistently or might
cause very high control costs. They are even worse by not having a limit where the growth
ceases.

c) Expansive species. It is often a very misleading increase pattern. For some reason some
species may exist in nature in low numbers forming just small populations or just few
individuals being scattered on a wide territory. This sleepy behaviour may end abruptly when
an opportunity created by human activity or by natural processes arises. A massive growth of
both populations and spread may occur in a very short time and in most of the cases there is
little to do to stop it other than by constant monitoring and eradication of severe infestations.
It takes usually a lot of time and great human and financial efforts to control such expansions.
Crassula multicava is one of the plants causing problems in several coastal areas; although
first noticed in 1957 and by 1959 considered by Healy to have a restricted distribution in the
South Island it is now widely accepted that its presence in nature may have occurred probably
around 1940 if not even earlier. As this plant was very popular in cultivation since the end of
the Victorian era I think we can assume without being too far off that it became adventive in
the early 1900s, had for decades just a sleepy behaviour without even being noticed only to
expand massively in the late 50s early 60s. Sometimes this expansion occurs just locally
Senecio serpens is a good example. Virtually unknown in New Zealand natural habitats
(except for a small population at Hunters Gully - Christchurch) this plant has caused few
years ago a severe infestation out of the blue on Somes Island (Wellington) luckily
eradicated. It is hard to say what actually triggers those expansions, it could be a combination
of environmental and climate factors involved, but also the capabilities of the species are very
important as well. This type of behaviour (if previously known) is very important for the
assessment of an alien plant; however, in a new environment plants habits may change
influenced by several other factors and become irrelevant a good example is the explosive
dispersal of some Opuntia species in Australia or Ethiopia, contrary to the normal and
balanced behaviour in their natural habitats.

d) Fluctuating species. The reason why plant populations may increase or decrease periodically
may be very complex and influenced by the continuous balance between habitats lost or
restricted for some reasons and natural dispersal or by the climate cycles; especially exposed
to the latter are the annual plants with populations increasing / decreasing depending on the
conditions but also few succulent perennials are affected. Usually the environmental agencies
are monitoring the populations during the peak years to prevent expansion but luckily most of
the naturalized species (probably over 80%) are fluctuating species. A. E. Esler 1988:
They are species whose populations changes are not evident or are considered to be short
term fluctuations. The reasons for low and seemingly low populations are seldom evident
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except where restricted habitat causes limitations, or measure such as weed control are
exerted against them. Some species have the potential and opportunity but have not time to
build up high populations. Others have low potential through lack of effective propagules,
lack of dispersal, or other causes.

e) Decreasing species. These are basically the species unable to adapt to the new habitats and to
form self-sustainable populations; the most unsuccessful of the aliens. Small groups or
scattered isolated plants may resist for some time in nature, sometime even few decades but
on the long run they have no chance of survival. Some of the adventive succulent plants
belong to this group. That this pattern is not determined by habitat destruction but mainly by
the inability to survive is proved by the fact that ca. 100 adventive species have disappeared
only from greater Auckland area over the last 100 years compared with only 5 endemic
species extinct New Zealand wide over the last 150 years.

It is always hard to include a plant in one or another of these categories as it is in some extent a
subjective judgment. The opportunities given are different and hazard plays in some cases a major
role too. The main thing is that alien species can find their way to success only if we allow it; in fact
we are paying now for the errors (read massive habitat destruction) done in the past. With the
increased attention given to our natural habitats over the last 50 60 years dramatic changes in New
Zealand nature are becoming more and more remote. It is also very important that New Zealand
habitats (a recent civilization less than 200 years) are far less altered by human activities compared
to Europe or even North America and therefore far more resistant to alien invaders. At least for
now.

However, there are also very important encouraging factors of which the most relevant are:

a) Habitat destruction. This is without any doubt the most important encouraging factor.
Habitat destruction may be a byproduct of human activities (urbanization, agriculture,
deforestation, public works, and earthworks of any kind) or a result of natural processes
(landslides, volcanic activities, floods, desertification, and erosion processes due to climatic
shifts) and can have a huge impact on the resistance to invasive plants. This phenomenon
may help create niches or pockets in which alien plants can establish populations and in worst
case can trigger irreversible habitat alterations, facilitating the massive dispersal of the alien
plants. Habitat destruction is the main concern because otherwise it is almost unthinkable that
succulent or xerophytic plants can establish significant populations in very compact, humid
and relatively dark habitats of the native bush. But the same plants can colonize easily
disturbed land as for example the roadside banks are. Sadly these are the main dispersal
corridors.

b) Natural niches. It is more or less applicable to the various coastal habitats, which are
somehow under populated by native succulents or xerophytes, except the salt marshes and the
sand dunes. The lack of native competition makes things easier for the alien plants; in
extreme situations like the recent volcanic island Rangitoto is is almost an open invitation
to colonize larger areas. However, in some cases plants like Carpobrotus edulis have
contributed massively to the stability and conservation of such habitats, particularly sand
dunes in this case.

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c) The lack of direct competitors or natural enemies. It is doubtless a very important factor
too, although I think it is generally overrated. It is favouring some species in particular
conditions but basically the expansion of the alien relies on the invasive character of that
particular plant and does not favour difficult species or of low potential. With or without
natural enemies or competitors most of the aliens never succeed. The damage done by
uncontrolled spread of few species (rather exceptions I would say) has contributed in giving
more weight, as it should be.

d) The presence of similar habitats or similar climate. Even in New Zealand pockets of
similar habitats or similar climate patterns corresponding to the needs of the alien plants are
not rare, especially for those of gondwanic origin.


VIII. Carpobrotus edulis, a friendly alien?


The Aizoaceae are well represented here in New Zealand and although most of them have just a
localized distribution it is nevertheless true that they thrive in some of the coastal habitats. The
species found here (4) are with just few exceptions originating from South Africa and inhabit mostly
coastal areas with a mild climate of Mediterranean type. The Mesembs are the next important plant
group after Crassulaceae, most of them original species but also few other cultivars. However, true
species or not over 15 different plants have been mentioned over the last decades. But the most
important for New Zealand and having a great impact on the sand dunes habitats of the coastal
regions are Carpobrotus edulis (Linnaeus) L. Bolus 1927, and (in a much lower extent) Carpobrotus
aequilaterus (Haworth) N. E. Brown 1928.



Carpobrotus edulis has actually a bad reputation as a very invasive plant, with a long list of places
where it had to be eradicated. But is it everywhere so? I am not arguing the extent of the
1. Sand dunes near the
southern end of Ninety Mile
Beach (Northland), where
extensive populations of
Carpobrotus edulis are
forming thick mats hindering
sand movement. Native
xerophytic grasses are also
present on big surfaces.
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environmental damages on other meridians and latitudes, but it seems to be a different story here in
New Zealand.



Basically there are two different ways in which Carpobrotus edulis has managed to naturalize. In
some countries (especially Europe, but also Australia and California) plants have escaped cultivation
and became invasive species posing serious ecological problems as they were competing indigenous
plants (some of them threatened or endangered) for resources, limiting their habitat and forming vast
monospecific areas. Of course every alien attempt to lower the biodiversity looks scary especially
nowadays, but we dont have to be that harsh in any given situation without a bit of reasoning. The
second mean of naturalization was in some cases (especially California and New Zealand, and
possibly Australia) that Carpobrotus edulis was deliberately planted to stabilize soil along railroad
tracks and later on was successfully used in stabilizing sand dunes habitats especially in coastal
areas. And thats exactly the point I want to make.

Carpobrotus edulis is originating from South Africa, growing on coastal and inland slopes and dunes
from Namaqualand in the Northern Cape through the Western Cape to the Eastern Cape, where it is
often seen as a pioneer in disturbed sites, but it became a quite cosmopolitan plant over the last
century or so. It is spread worldwide in areas with Mediterranean type of climate, usually in coastal
areas, becoming sometimes a pest and endangering the native flora. By only reading the common
names of this plant you will get a highly true image of its dispersal. Carla D'Antonio (2006):
Common names: balsamo (Catalan-Spain), Cape fig (English-South Africa), figue marine (French),
freeway iceplant (English-USA), ghaukum (Afrikaans-South Africa), ghoenavy (Afrikaans-South
Africa), highway ice plant (English-USA), higo del Cabo (Spanish), higo marino (Spanish),
Hottentosvy (Afrikaans-South Africa), hottentot fig (English-USA), Hottentottenfeige (German),
iceplant (English-New Zealand), ikhambi-lamabulawo (Zulu-South Africa), Kaapsevy (Afrikaans-
South Africa), patata frita (Catalan-Spain), perdevy (Afrikaans-South Africa), pigface (English-
Australia), rankvy (Afrikaans-South Africa), sea fig (English-USA), sour fig (English-South Africa),
suurvy (Afrikaans-South Africa), umgongozi (Zulu-South Africa), vyerank (Afrikaans-South Africa).
And further down we have the confirmation: Known introduced range: Portugal, Italy, France,
2. On Whangamata Beach
(Coromandel) Carpobrotus
edulis (the yellow flowering
form) is present just on smaller
patches, growing among native
succulent sand dunes flora.
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United Kingdom, Ireland, Germany, Spain, Canary Islands, Balearic Islands, Gibraltar, Malta,
Albania, Greece, New Zealand, western USA, Australia, St Helena, French Polynesia, Pitcairn,
Mexico, Chile, Argentina, Tunisia. It is definitely not an exhaustive list as Turkey and Hawaii are
missing.



It may be a very invasive plant with highly destructive potential for well-established native habitats
worldwide, but I want to point out that at least for New Zealand the benefits of the integration of
this plant in the coastal habitats are exceeding the downsides of the environmental pressure it puts on
native and endemic plant species.


3. Leaves turn red when
very strong light conditions
meet a marked water deficit
yellow flowering form on
Mututapu Island (Hauraki
Gulf).
4. Flowers of the pink
flowering form of
Carpobrotus edulis, photo
from 2004, unknown
location. (Photo by Carsten
Niehaus)
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Carpobrotus edulis was introduced in New Zealand by man some 150 160 years ago as garden
plant but apparently escaped very quickly in nature, and spreading quicker than one would think,
becoming in time a part of the local ecologic systems, and fully integrated in the New Zealand
marine landscapes. At some stage it was used also to stabilize sandy soils and sand dunes in coastal
areas. Carpobrotus edulis is quite common in many places. It is a mat-forming succulent which can
have a highly destructive impact on invaded habitats by smothering the existent vegetation and
reducing the natural regeneration the native flora and if thats not bad enough it may drive
changes to soil pH and nutrients components which could lead potentially to tertiary successions; at
this point it will be even harder or even impossible for the native flora to recover without human
assistance.







What makes New Zealand different, why isnt it here perceived as a serious threat for the native
coastal flora? I think you have to see and evaluate carefully both sides of the story.

5. New yellow flower from Motutapu Island (Hauraki Gulf). Flowers open several days in a row,
in some clones the yellow flowers change colour (becoming pinkish or whitish), as they get older.
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To start with and this is very important - the coastal areas of New Zealand are extremely varied (5),
from sandy beaches doubled with sand dunes, rocky outcrops, platforms and cliffs, followed by
rocky plateaus and ranges, with sometimes a quite alpine aspect just few hundreds of meters from the
waterline, numerous lagoons, estuary and fjords, offering an unbelievable wide range of habitats and
niches in which adventive vegetation (not necessarily succulent) can form stable and well settled
populations.



First of all Carpobrotus edulis does not limit in any way the natural habitat of an iconic endemic
the Maori Ice Plant or Horokaka, Disphyma australe ssp. australe by its Latin name. The two
although similar in some extent and both belonging to the Aizoaceae family have quite different
preferences; Disphyma australe ssp. australe is more often seen in rocky habitats, rocky outcrops or
even vertical cliffs and is confined to the waterline in most of the cases. On the other hand
Carpobrotus edulis prefers sandy banks or a typical sand dune habitat, being able to penetrate the
inland along railroad tracks or roadsides though, probably similar to the reported USA populations
(6); it is definitely not strictly confined to coastal areas although it seems to prefer marine exposure
and the good natural ventilation that comes with it. Both species come along together very well and
there are also several sites where both species occurs. There is also a very interesting (to say the
least) natural hybrid, a unique and surprising enrichment of the biologic genes pool of this kind of
habitats I would say. I wrote about this natural hybrid in an earlier article (7).

It is very important to understand that some of the coastal habitats have a very fragile nature; salt
marshes and sand dunes for example are probably the most exposed to weathering or erosion. A
slight change of currents (for salt marshes) or strong storms with battering winds (for sand dunes)
can dramatically change the environmental factors dunes are relocated, shores are reshaped and
lagoons are appearing or disappearing. There are definitely more chances of stability when well
settled vegetation occurs and less damage is done by weathering or erosion. Carpobrotus edulis was
introduced in New Zealand possibly 150 160 years ago and used shortly after that for erosion
control especially in unstable soils or sand dunes near farmland. Later on it has been successfully
6. A flower from
the same Motutapu
Island population,
several days old,
already turned
almost white.
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used for erosion control and dune stabilization in coastal area. One of the many places where it was
deliberately used for this purpose is Long Bay, on the east coast of Auckland where I have seen it in
large numbers and on reasonable big areas only few years ago (8).








However, at some stage, after the European colonization took off, the use of Carpobrotus edulis for
soil stabilization would have been definitely the better option. Not only are the beach and sand dunes
habitats extremely exposed to erosion but also they are a highly important support for seabird
populations as nesting habitat. There is no doubt that the expansion and invasion of Carpobrotus
edulis has put the native and endemic flora under pressure but lets face the truth it could have been
worse than that: completely unstable dunes and fore dunes with no vegetation at all, as we can see on
the northern end of Ninety Mile Beach, at Te Paki (where boogie boarding is a great fun for tourists),
huge loose sand dunes with Saharan aspect. Coastal habitats are typically ideal for plants with
xerophytic adaptation (not necessarily succulent plants) as the substrate is mostly very shallow and
permeable or even quite loose, unable to retain moisture for too long. In coastal habitats substrates
are therefore mostly dry within few days and additional encouraging factors are usually exposed sites
with strong winds, harsh sun and in some regions short periods of draught in summer this means
xerophytic plants do usually better. It seems to be a paradox, but in fact it is not not too long ago
New Zealand climate was sensibly cooler and rather on the moist side; this explains (at least partly)
why the coastal areas of New Zealand is rather under-populated with plants adapted to environments
encountering often a water deficit, suggesting that an ever-wet climate has been replaced by one
where there was at least seasonal dryness. Even in areas with commonly higher rainfall levels (9) the
coastal habitats are largely under-populated with native plants having xerophytic adaptations
(including here the succulent plants) or any other plants. Arguably with the exception of the sand
dunes there is no serious competition for alien plants in coastal habitats of the drier type and there is
no wonder that invasive alien plants are taking all opportunities. In fact even the most settled and
unaltered coastal habitats have a higher density of naturalized and adventive flora compared to bush
lands or alpine habitats.

7. 8. Spreading mats of yellow flowering forms of Carpobrotus
edulis at Tahuna Torea (Auckland) on the left and Motutapu Island
(Hauraki Gulf) on the right. As you can see in both photos collected
debris may hinder the establishing of other xerophytes.
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In a compact and settled habitat Carpobrotus edulis would have had less chances of intrusion as it
mostly seen as a pioneer in disturbed areas. Beside the usually low plant density, sand dunes habitats
are often naturally disturbed and in some degree the soil instability is rather benefic. Given the
opportunistic behavior of xerophytic plants, these can conquer and settle newly disturbed areas
offering well-drained substrate, plenty of sun and room to spread. These cycles of settling /
disturbances are in fact characteristic for dunes and fore dunes. Carpobrotus edulis may hinder this
natural process as it usually forms very compact and dense mats, 15 30 cm deep and up to few
meters wide, not only smothering existing vegetation but breaking this natural disturbance cycle.
Unless heavy frost damages occur or nitrogen soil levels naturally increases there is little to stop it
from spreading.

It may become a nuisance in time (to say the least) but it is not that easy to get rid of it. By simply
removing the plant you just create a disturbed space for open competition, and as most of the time
happens, it is not for the good of the native flora. Tertiary plant invaders, already present in small
numbers or having dormant seed banks waiting for the right conditions, can take advantage of this
and may spread uncontrolled and benefit from the accumulation of nutrients facilitated by
Carpobrotus edulis. This all could be for the worse as it actually impedes restoration processes. To
be effective the removal of Carpobrotus edulis has to be followed by substrate conditioning if
necessary and adequate planting of native / endemic vegetation. Also a constant monitoring for at
least 6 12 months is necessary. The restoration of Long Bay dunes mentioned earlier has probably
followed these steps, as it seemed to be quite efficient. Only few iconic patches of Carpobrotus
edulis were maintained. Carla D'Antonio (2006) maintains that for best results Carpobrotus edulis
should be initially selectively removed in order to allow rapid soil stabilization and minimizes sand
movement in the area; later on, after the area has been successfully restored to form strong native
plants communities, the remaining Carpobrotus edulis can be removed.

9. Carpobrotus edulis
does very well in pure
fine sand the pink
flowering form at Ninety
Mile Beach (Northland).
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Widely promoted only few decades ago, successfully used for erosion control in the past, this plant is
still a part of the New Zealand ecosystems, fully integrated in its natural marine landscapes. Its a
statement that alien plants can be useful at times and that before judging mans interference with the
course of nature one has to see both sides of the story.



Carpobrotus edulis is widely spread in New Zealand and forms in some places strong populations.
Extended populations can be seen throughout Ninety Mile Beach in Northland, on the western
Tasman Sea coast, which is probably the most fabulous sand dunes habitat in New Zealand. I have
seen smaller patches in largely different places such as Long Bay (Auckland, on the East Coast),
Motutapu Island and Rangitoto Island (Hauraki Gulf), Tahuna Torea (Auckland), Muriwai (just out
of Auckland, on the West Coast), Whangamata (Coromandel Peninsula), to name just a few. I have
collected adventive succulent plants from natural habitats on occasion as I think that they really dont
belong here in New Zealand despite the interest I have for plant naturalization, but I have never
collected Carpobrotus edulis as this is my opinion it has earned its place in natural habitats and is
now fully integrated.

Additional References:


Carla D'Antonio (Professor Ecology, Evolution & Marine Biology University of California, Santa Barbara USA) - Carpobrotus edulis (2006, ex Global
Invasive Species Database http://www.issg.org/database);

Dr. Curtis Daehler - A Risk Assessment of Carpobrotus edulis for Hawaii and other Pacific Islands (2006);

Christien Malan & Alice Notten (Kirstenbosch National Botanical Garden) Carpobrotus edulis (2006, ex www.plantzafrica.com);

Didge Rowe Cultivating Mesembryanthema in New Zealand (New Zealand Cactus and Succulent Journal, Vol. 59 - 61, 2006 - 2008);

Wikipedia, the free encyclopedia (English version).

10. Tahuna Torea
(Auckland) at low tide,
with small patches of
yellow flowering form
of Carpobrotus edulis.
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Page 14
Further Readings:

Mesembs of the World (1998) a comprehensive monograph edited by the National Botanical Institute of South Africa.

My Notes:

(1) It was not observed seed forming in New Zealand plants but Crassula multicava (ssp. multicava as Tlken considers) has tiny plantlets borne in
the axils of the inflorescences which are easily dispersed by wind.

(2) As my referred readings and my own observations are strictly limited to the New Zealand natural habitats it is to understand that not everything
might be applicable or have the same importance in other parts of the world. On the other hand it is not my intention to make an exhaustive account on
this subject.

(3) From the 300,000+ plants and horticultural hybrids considered to have been cultivated in New Zealand over the time less than 10,000 have been
ever observed in a natural or semi-natural environments, less than 2,600 have been considered additional, casual or naturalized at any time, less than
800 are considered now naturalized (being able to self-maintain populations), and less than 300 are a real threat for New Zealand nature. This might not
be true anywhere else, but it is here in New Zealand, at least for now.

(4) Aptenia, Carpobrotus, Delosperma, Dorotheanthus, Drosanthemum, Disphyma, Lampranthus, Oscularia and Ruschia it is quite a list!

(5) It is also very important I think to state that in an insular territory like New Zealand the coastal regions have a greater overall importance in its
biologic profile compared with continental units; except for New Zealand (and very few other territories like Hawaii) Carpobrotus edulis is spread only
on coastal regions of more or less continental blocks. This brings (and I think there is no surprise in it) a different set of pressure factors on the native
coastal flora in play, an additional enemy such the invasive Carpobrotus edulis having definitely a greater impact.

(6) Not at elevations higher than 150 m though.

(7) The first article of the series dedicated to the New Zealand succulent flora Succulent plants from own under Disphyma australe and its natural
hybrids (2007).

(8) It was probably in early 2003 when I saw on large surfaces growing this plant (at that time mistaken by me for Lampranthus aurantiacus);
nowadays large patches have been removed and replaced by native sand dunes grasses and plants, just couple of larger spots remaining by early 2007.

(9) For more details on New Zealand climate you can consult the chapter Origin, Climate and Coastal Habitats in Succulent plants from own under
Adventive Plants (Part 2) (2007)


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All errors, omissions and misconceptions are mine.

All photos by Eduart Zimer (except no. 4 by Carsten Niehaus, ex Wikipedia, the free encyclopedia)




Eduart Zimer, November December 2007 & June - July 2008

http://eduart.page.tl/

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