Eduart Zimer - (SDU) - Adventive Plants - Part 4 (2008)

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Succulent plants from down under Adventive Plants (Part 4):

Natures Laboratory: Rangitoto Island (Hauraki Gulf) A brief overview of the adventive
succulent flora of Rangitoto Island

IX. Natures Laboratory: Rangitoto Island (Hauraki Gulf)

The volcanic island of Rangitoto (Hauraki Gulf) is always a very interesting subject not to mention its
abundant succulent naturalized flora. The island is an extremely recent appearance at geological scale and
is therefore an excellent opportunity to study the appearance and development of flora and fauna in an
environment dominated until not too long ago by bare lava fields (1). It is in other words a natural
laboratory. Geologists say the island emerged some 600 800 years ago from the sea, although there is no
generally accepted opinion, following a series of violent eruptions lasting with interruptions between 10
200 years (depending on who is accounting). However, at the time of the main wave of the Maori
settlement (probably between 1250 and 1300 AD, not later as wrongfully still common considered), the
volcano was active and sending its ashes high in the sky. Even after lava eruptions have ceased it is
believed that there still was a significant volcanic activity (gases, tephra eruptions) for several centuries
until shortly after Cooks voyages possibly until 1780 1790.

Rangitoto wasnt really inhabited by Maori tribes, although they must have spent some time there as there
are signs of their presence on the island, but the neighboring Motutapu Island was. Later on there was a
small European community living on the island, before becoming a sort of holiday baches community. At
some stage (until shortly after World War II) there was also a prison on the island providing of course
cheap labour for a small quarry; however, no major economic activities have been started here and
1. Photographs and
drawings from the 19
th
century prove that the
vegetation was far from
being established at that
time. This ink and
watercolour painting by
Charles Heaphy shows the
volcanic cone and the
crater of Rangitoto in the
1850s. It is believed that
in those times the cone
was bare of vegetation,
largely because of fires lit
by early European
settlers, but I really doubt
this. The desolated
landscape reminds more
of recent volcanic activity.
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fortunately there was little ecologic impact. At any point in time (if we exclude the residents of the prison)
there was no significant permanent population living on the island, probably not exceeding 50 people.
Probably the main reason for such unimportant settlements on the island was the lack of fresh water. The
few residents had a vivid lifestyle though, a source of colourful memories for many families, including
fabulous contests and games, a legendary swimming pool and memorable events like the day the southern
wharf collapsed in 1955, nearly killing few officials. By 1915 two fine gentlemen named Mr. Leary and
Mr. Wilson even started the works for a Botanic Garden, including reportedly many cacti and other
succulents, never finished and eventually abandoned as it has happened with many other projects during
troubled times of war. But by the end of the 50-ties the quarry and the prison were already a thing of the
past and most of the baches were abandoned as strict rules were set in place.

At present no economic activity whatsoever is allowed on Rangitoto Island, except the two trucks hauling
tourists to the summit or for runaround trips. Even the tracks are largely ignoring the northern half of the
island, except for the path leading to Wreck Bay and Boulder Bay. Wreck Bay used to be until late in the
40-ties a place where ships were set ashore or deliberately sunk in shallow waters and abandoned after the
service days were over it is a graveyard of vessels simply dumped on the island between 1887 and 1947.
2. The crater of Rangitoto in 2006 quite dense vegetation consisting of Metrosideros excelsa
(the iconic pohutukawa), Metrosideros robusta, Erica lusitanica and Hypericum androsaeum.
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No less than 13 ships are resting now mostly underwater, being just an attraction for tourists and especially
for divers.

Rangitoto Island is very interesting for botanists, biologists and ecologists because they can watch in real
time the entire process of colonization of the lava fields, first by lichens and after some time by vascular
plants which will continue to change the environment until bush land and forest-like habitats can establish
themselves. This process of development and diversification of the vegetation allows land animals and
birds to settle and to form significant populations. It is a process, which can be monitored nowadays in just
very few places on Earth, and therefore it is understandable why scientist whish no interference with these
natural processes. It is not always easy as animals (goats, bewildered dogs and cats, possums, wallabies,
rabbits, hedgehogs, mice, rats and other mammals) were affecting the natural processes by destroying (in a
very selective manner sometimes) the young shoots of some of the plants, foraging widely on the ground,
eating ferns or defoliating and depleting shrubs, destroying seedlings or hindering and endangering the few
bird colonies, changing massively the unsettled natural habitats. Of course all this was disturbing the ways
of the nature and the extent of the damages done have forced scientists into drastic measures like more or
less brutal eradication of some of these animals or at least for a drastic control of their numbers, for the
despair of and accompanied by the protests of many animal rights associations. During 1990 2000 years
this matter was almost settled but the numbers of loose mammalian browsers is still monitored and
controlled.

There are also many blacklisted intruders, some of them naturalized succulent plants. There are
eradication and control programs in place; each species was carefully evaluated and monitored and as a
result there is a target set Eradication, Zero Density (2) or Sustained Control. However, it is remarkable
how many alien species (including many succulent plants) have been naturalized on the island.

One may ask few legitimate questions why are that many introduced plant species on the island for
instance, even more species than in adjacent Great Auckland area or in other centers of early European
settlement, the typically introduction areas? Why are the succulent intruders especially well represented on
the island? Why do we have to find here on Rangitoto succulent adventive flora not even present in other
areas where these plants are heavily cultivated?

Compared to the mainland or even other island habitats Rangitoto plays definitely by some other set of
rules. The main centers of pressure for the natural habitats were the oldest areas of European colonization.
From here has started everything for the classics of plant naturalization, the escape and initial dispersal -
Paihia and Russell (Bay of Islands in Northland), Auckland, Waikato, Bay of Plenty, Canterbury,
Christchurch and Otago (3). Deforestation, extensive farming, the rapid extension of pasture lands, the
introduction of new plants and grasses from the old world for agricultural purposes, not to speak of the
cottage gardens packed with all kind plants of horticultural origin or of the large number of abandoned
settlements (during or immediately after the gold rush or even later as a byproduct of urbanization) all
these had a large impact on habitats and as a result on the number of the naturalized plant species, but in
none of these regions the relative number of intruders reported to the total number of species was so high
as it is in Rangitoto, not even close, although human interference with natures business was of
incomparable lower extent here.

Rangitoto Island is definitely the most dynamic habitat in New Zealand, evolving in a fast pace. In such an
unsettled environment, largely still unbalanced, each and every new species introduced can have a
tremendous impact and may represent a potential danger as it can easily find from the multitude of existing
ecologic niches a suitable one. A good proof for that is the neighboring Motutapu Island separated by some
20 to 200 meters of shallow waters, a distance which can be crossed by foot during low tide. Much older
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and with totally different geologic origin, Motutapu Island has an incredible different flora and fauna,
actually very visible even if you look at them from the mainland. Although most of the means and patterns
of plant colonization have been already mentioned in this series I will reinforce briefly few of the factors
contributing to a significant presence of adventive succulent flora on the island, the ones I think have more
weight in this ever-changing habitat:

the lack of deep rich organic soils is not a barrier, not even a handicap for succulent plants.
In the back of our minds we bear the idea of rich organic soils necessary for plants in order to
thrive, we probably are highly influenced by an agricultural or horticultural way of thinking
get them plenty of organic matter and fertilizers on top of that and you will have a rich
harvest. In the real world of the natural habitats plants seldom need very rich soils, most of
them do it very well with moderate to poor soils. Especially the succulent plants prefer mostly
poor to mineral soils. Despite their rather slow growth and dispersal succulent plants managed
to cover this handicap by a differential gap most of them can grow without any soil at all!
Very briefly (4) most of the plants cannot grow on bare rocks; therefore lichens are first
colonizing the lava fields. After some time they will decay and produce small amounts of
organic matter, accumulated eventually in small pockets, fissures or rifts first step of the
colonization process. Eventually these accumulations of organic matter will retain enough
moisture to allow seeds dispersed by wind or birds to germinate and start the actual colonization
process of higher vascular plants. From now on the snowball effect is in place, more and more
decayed organic matter will be produced, changing the environment and forming shallow soils
with organic content, rich and deep enough to support bush vegetation, giving moisture
retention another boost by a lesser direct sun exposure of the ground, which especially in high
rainfall areas (as Rangitoto is) creates at least the minimal conditions for a future rainforest. All
this takes many decades or most likely few centuries. Most of the succulent plants dont need
all this process, they can accommodate themselves on shallow mineral soils (as a result of
erosion, although that is not usually the case in such a young volcanic island, or more likely
accumulations of volcanic ashes from the previous eruptions) or with roots cramped directly on
the lava blocks, exploiting rather the richness in vital minerals. As all human settlements on
Rangitoto were in the proximity of the shores, these were the most exposed areas where
3. 4. The gap between Rangitoto and Motutapu at the narrowest point, between Islington
Bay and Gardiners Gap, at low tide (left) and shortly after the tide started to rise (right).
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succulent plants (and other plants with xerophytic adaptation as well) have been dispersed, not
the inland that took the rather traditional path of the development to a light bush. With very
few exceptions the adventive succulent flora of Rangitoto is limited to a narrow coastal strip
following mostly the inhabited parts of the island.

the scarcity of available water is another reason why the succulent and xerophytic flora had a
significant advantage on more moisture needy species. Unlike the neighboring Motutapu Island
there are no fresh water sources on Rangitoto and thats a serious handicap. Even if the rainfalls
are quite abundant in the area (averaging 1,100 1,300 mm / year in Auckland and although
there is no meteorological station on the island it is believed that the rainfall is marginal higher
here (5)) the water flows simply away as there is no deep soil to retain it. Some moisture will be
retained in deeper pockets of the lava fields or elsewhere where soil has started to accumulate
but this is hardly enough for plants to survive, therefore plants with xerophytic adaptation
(including succulent plants) are definitely better off, at least until the environment changes
enough for a secondary succession. The scarcity of water is a very important factor not only
because it limits the range of plants, which may establish here, but it also affects the structure of
the fauna - which in the end supports plants dispersal or influences plant successions.

the absence of any competition is a very important factor as well, as alien plants find in most
of the cases very difficult to penetrate and settle in very established habitats reigned by different
environmental parameters. However, it is amazing that in a relative short period of time few
hundreds of plant species have established here, including 40 species of ferns and ca. 25 species
of different succulent plants. There is definitely much more variety than one would have
expected. Photographs and paintings from mid to late 19
th
century show a far less populated
island, with large areas of bare lava fields. Many species including succulent plants have found
the right conditions, plenty of space and a variety of available niches, good enough for these
opportunistic intruders.

5. Bare lava fields, this is
how all starts. But the
very most of the succulent
plants manage to
accommodate themselves
in this hostile
environment, exploiting
the richness in vital
minerals of the rocks. On
the contrary, the common
vascular plants are
waiting first for the
lichens colonization to
transform the habitat.
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the human interference was more effective than in other given circumstances. There were
several attempts during the late 19
th
and early 20
th
century to beauty the harsh volcanic
landscape, desolated and maybe depressing for the few inhabitants. Even planting days were
organized in the 1890s when not only the inhabitants but also members of the public were
encouraged to bring by ferry their favourite plant with them, not to mention Mr. Learys and
Mr. Wilsons botanical project which lead to the introducing of two of the most widely
established plants in the 259 m high summit cone area Erica lusitanica (fam. Ericaceae) and
Hypericum androsaeum (fam. Hypericaceae). Approximately 60% of the plant species present
on the island, some of them considered now to be invasive on Rangitoto and potentially
disturbing and re-phasing natural processes, were introduced during this period mainly as
garden plants. The very most of the species which succeeded to escape and naturalize were
succulent plants (Crassula, Sedum, Bryophyllum, Aloe, Aeonium, Agave spp., etc.) or plants
having xerophyte adaptations and features like bulbs, rhizomes or corms, enabling them to
survive in extreme dry environments (Watsonia, Gladiolus, Iris, Nephrolepsis, Asparagus spp.
and many others). The human assisted dispersal was far more effective than natural dispersal
means like seeds dispersed by wind or by birds for example, accounting for only 40% of the
plant species.

It is not only human interference diverting natural processes and setting a mark on Rangitoto natural
history. It is also rather strange though that the vegetation development patterns are not closely
following the typical New Zealand vegetation types (or what one would have expected to happen in
New Zealand) but rather Hawaiian like forests or light bushes dominated by trees belonging to the
genus Metrosideros (fam. Myrtaceae), especially by Metrosideros excelsa (the famous native
pohutukawa, one of New Zealands iconic plants) and Metrosideros robusta (Wotherspoon &
Wotherpoon, 2002). A hybrid swarm of the two was noticed but it is thought that it is progressively
backcrossing to Metrosideros excelsa. It looks rather odd, but Rangitoto Island has the largest
pohutukawa population known in New Zealand. This iconic tree used to cover large areas and form
massive forests, but it is becoming increasingly rare on the mainland. Another remarkable thing is that
6. Like a stairway to
heaven, these steps lead to
the heart of the young bush.
Such artifacts are often seen
in the proximity of the shore
in Islington Bay and
Rangitoto wharf. The bad
news is the men have
already interfered too much
with the natural flow of life
in all forms here on
Rangitoto. The good news is
that nature has a
tremendous regeneration
power.
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the layer ground of the island, the hostile lava fields, is the main support for many plant species which
in the mainland forests grow as epiphytes for example Collospermum hastatum and Astelia banksii
(both belonging to fam. Liliaceae) and Griselinia lucida (fam. Cornaceae) which makes no sense at all
as the soil has little or no water retention at all and is totally different from a classic epiphyte substrate.
Fact is that many processes are rather different from the mainland patterns and this might be another
quite sustained explanation for the high occurrence of succulent plants from the total number of
species.

Most of the common available sources consider that there are 200 230 different plant species on the
island (6) but it looks like this number is largely understated. Wotherspoon & Wotherspoon, 2002 (ex
Gardner, 1997) are maintaining that there are at least 232 naturalized vascular plant species and 286
native vascular plant species on the island, giving Rangitoto flora a composite structure and a quite
unusual weed population. Luckily the very most of them have little to no impact, some are quite a
nuisance at times, and some of them are posing a real threat due to their pronounced invasive character.

X. A brief overview of the adventive succulent flora of Rangitoto Island

It is quite hard to get a full picture of the adventive succulent flora of Rangitoto as many inaccuracies and
confusions have been made in time, and I might add some as well. However, apart from the fact that we
have to define the term succulent plant which draws right from the start the intended limits of observation,
7. Light bush on Boulder Bay track. This is the typical vegetation status on the island.
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I am relying mostly on written botanical accounts and few environmental studies, but also on personal
observations made during my two trips here. It is understood that the following list of names is not
intended to be an exhaustive account but rather to bring some light in an area which was until now largely
ignored as a whole.

By succulent plant I understand any perennial vascular plant having evolved specialized organs for the
storage of water and nutrients (like caudexes, trunks, stems, and leaves) as a form of xerophytic adaptation
in order to allow them for many years the survival in an environment characterized by a marked seasonal
or periodic moisture deficit. I am therefore excluding annual plants (mainly concerned to consume the
reproduction cycle in a given period of time), or using succulent tissue in order to increase the survival
chances of their descendents (like plants having berry-like fruits), or being able to survive prolonged
periods of draught due to their bulbs and rhizomes even if parts of the plant die back. It is certainly not a
proper definition of succulent plants, and probably will raise some eye brows (as it did in some other
occasions), but I think it defines better my area of interest and the intended limits of my account.

And here we go:

Agave americana Linnaeus 1753 (fam. Agavaceae) [Mexico, USA] is one of the plants never mentioned
in botanical accounts (but in some of the eradication programs Wotherspoon & Wotherspoon 2002, not
in Craig J. Miller 1994), although its presence in other Hauraki Gulf islands was acknowledged. It is one of
the earliest botanical accounts on naturalized succulent plants, mentioned in 1969 by T. Kirk as growing on
Aucklands North Shore. I have seen the plant on the eastern coast of Rangitoto, just north of the Islington
Bay wharf, forming quite abundant populations of rather young plants, none larger than 100 cm, scattered
over fairly open but very rough lava fields. I havent seen any large adult plant, nor variegated variants as
reportedly seen in other places (7). The short-term goal of the Department of Conservation is to achieve
8. Lots of Agave
Americana growing
on very shallow
rocky soils (ashes
from the terminal
phases of the
volcanic eruptions)
or directly on the
lava fields. This
reminds me on the
Agave americana
variegata seen by me
in 2003 growing in
pure fine sand in
Mount Maunganui;
what a difference!
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Zero Density (apparently already achieved) the long-term goal being the full Eradication it is a Class 2
priority plant. In this case the problem in achieving Zero Density is that even younger plants are able for
vegetative reproduction due to their rhizomes, and in this case they surely do so at least three or four as
many just few centimeters small plants were scattered around them. I have collected three 5 6 cm large
plantlets for further observation. My impression is that the Rangitoto population belongs to the form
usually known as var. protoamericana, but I might be wrong.

Carpobrotus edulis (Linnaeus) L. Bolus 1927 (fam. Aizoaceae) [South Africa] is reportedly quite
common on the island, although I have not seen it growing on Rangitoto, but on the neighboring Motutapu,
very close to the narrow passage between the islands. I wrote extensively in Part 3 of this series about this
plant so that I will not insist. Apparently just the yellow flowering form is present. It is another Class 3
priority plant, although I see no reason why it shouldnt be left alone. However, all efforts to achieve Zero
Density make no sense at all as it is a fast growing plant, usually flowering and setting seed within 1 year.

9. 10. Agave americana, smaller plants growing in
abundance south of the passage to Motutapu Island.
11. 12. Carpobrotus edulis grows
abundantly on Rangitoto, but not
on the eastern coast where I was
browsing during my second trip
here. This plant exposed to high sun
levels was photographed less than
50 m from the passage to Motutapu,
but on Motutapu Island (left). Big
patches of the yellow flowering
form (below on next page) were
covering the low sandy soil of the
beach.

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Aloe maculata (Aiton) Haworth (fam. Asphodelaceae) [South Africa] referred mostly as Aloe saponaria
(Aiton) Haworth 1804 in New Zealand botanical records, is a very popular and easy-going plant, and there
is no wonder that it managed to settle here. It was firstly mentioned in 1883 by T. F. Cheeseman as Aloe
latifolia (Haworth) Haworth (8) in Remuera (Auckland). It is a quite slow growing plant especially in harsh
conditions and with a rather low dispersal rate. Rhizomes are sent around the mother plant and eventually a
medium sized compact colony is formed. Contrary to cultivated plants or wild plants growing in different
conditions (as I saw in July 2003 in Wenderholm, north of Auckland, in a light forest at 150 - 200 meters
from the sea) Rangitoto plants remain rather low growing, in semi-sheltered positions on lava fields. It
might have been the intense heat at ground level, or the lack of water the plants I have seen were looking
rather stressed and most of them of brownish colour. Of course the plants were mostly forming small
groups around older plants (or around already dead plants) but without forming dense colonies. The plants
I have seen were scattered on the eastern coast of the island, south of Islington Bay wharf, close to the sea,
in few distinct populations. It is quite possibly that the plant was cultivated in gardens and that originally it
was a typical garden escape; however, as I havent seen any plants growing close to abandoned baches, or
their former gardens, I am pointing rather to seed dispersal by other means, at least for the newer
generations. It is a Class 3 priority plant short-term goal being Sustained Control and long-term goal Zero
Density. Overall it is a very colourful addition to Rangitoto flora.

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The rest of the adventive succulent plants on Rangitoto Island belong to Crassulaceae, which is no surprise
at all as this plant family became increasingly popular in New Zealand gardens and greenhouses over the
last century. I will start with a very interesting (and disputed) group of plants forming sensational
populations Aeonium.

I will start with a plant having an uncertain status Aeonium arboreum Webb & Berthelot [Morocco]; it
is unknown if this plant has been actually seen on Rangitoto or anywhere else in New Zealand although
there are a few references. Apparently it was Healys confusion in 1959 (reportedly he confused Aeonium
cf. ciliatum for Aeonium arboreum), many have referred him later, but when his error has been discovered
years later all authors tend now to link any reference of Aeonium arboreum to this error, which might be of
course not true. However, little is actually known other than this plant is a doubtful occurrence.

Aeonium ciliatum (Willdenow) Webb & Berthelot [Macaronesia] is one of the succulent plants early
mentioned by Healy (1959) and acknowledged in 1984 by D. R. Given as being widely dispersed in both
North Island and South Island although not forming dense populations.
13. 15. Mature Aloe maculata (above
left) growing directly on the lava fields.
Note the brownish colour showing
intense stress (intense sun, constant
deficit of water or both) and the rather
thin long leaves usually plants
exposed to strong light levels have
shorter leaves but broader at the base.
Stressed young plants growing directly
on lava blocks (above right) and on
shallow rocky soils (below left). The
plants seen here are quite different
from the small colony seen in 2003 in
Wenderholm in a sheltered position in
light woodland, reminding me more to
cultivated plants.
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Aeonium cf. ciliatum (Willdenow) Webb & Berthelot 1841 [Macaronesia] just few isolated plants not
matching exactly the descriptions of the original species; it is different from Aeonium ciliatum mainly by
having the leaves fully green (and not with a red border in strong light conditions) and by its greenish
flowers (opposed to reddish in Aeonium ciliatum). Some authors consider this plant a hybrid with Aeonium
urbicum and it is also possibly that A. J. Healy has confused this variant in his 1959 account, an error
perpetuated later also with other populations, with Aeonium arboreum. D. R. Given mentioned it for the
first time in his 1984 Checklist.

Aeonium x floribundum Berger [hybrid of garden origin] apparently a single well settled population
very close to the high tide mark. This hybrid was mentioned as early as 1988 by W. R. Sykes in vol. 4 of
Flora of New Zealand, based on a 1986 collection. Between 1989 and 1993 Sykes has collected this
hybrid from Hauraki Gulf islands (published by P. B. Heenan in his 1998 Checklist), including Rangitoto.

16. A wonderful specimen of
Aeonium undulatum hybrid
growing just south of
Islington Bay wharf. There is
a magnificent strip of
Aeonium hybrids growing
close to the water line
cramped on the lava block.
This Aeonium population was
definitely the highlight of the
second trip to Rangitoto,
probably the most spectacular
succulent scene I have seen in
the wild.
17. Aeonium ciliatum (possibly a
hybrid plant) growing in the same
location. It is amazing how
different the plants can be; it is next
to impossible to find two identical
plants. The plants here have
definitely not the horticultural
look we are used to. However, even
if botanic literature states that only
isolated plants were seen, I think
there were at least 30 40 plants I
have seen on a 100 150 m long
strip. Very few young plants
though.
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Aeonium haworthii Webb & Berthelot 1841 [Canary Islands] mostly hybrid plants have been collected
from several places in New Zealand, but none of the botanical literature references is placing it in
Rangitoto. However, it is present on the island and mentioned in Wotherspoon & Wotherspoon (2002) as a
Class 3 priority plant targeted for Zero Density on the long run, not mentioned in Craig J. Miller (1994).

18. In the same
location Aeonium
undulatum (left) and
Aeonium ciliatum
(right) growing next to
each other of course
both are hybrid plants,
but at least one of the
parents is clearly
visible.
19. A wonderful
rosette of an Aeonium
haworthii (a hybrid
plant I think). This
specimen is probably
one of the most
resembling the true
species; there were
some other as well,
but also many
indefinite forms.
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Aeonium undulatum Webb & Berthelot 1841 [Canary Islands] - few isolated plants growing close to the
tide line. First collection dates from 1986 (W. R. Sykes - published in vol. 4 of Flora of New Zealand in
1988) and was made in Rangitoto, other populations being discovered shortly in Banks Peninsula
(Christchurch). The plant is mentioned in botanical literature, but not in environmental studies or
conservation programs for a change.

20. Another clump of
Aeonium haworthii,
quite common in the
area and apparently
being established for a
long time, as older
specimens were also
available. It is rather
odd that botanical
literature does not
mention this species
(or its hybrids) here
on Rangitoto Island.
21. A view with a quite
surrealistic touch of
the Aeonium colony
growing on lava
blocks close to the
high tide line and
definitely having their
share of seawater mist
during storms. Mixed
form of Aeonium
undulatum and
Aeonium ciliaris are
good visible, note also
the many plants which
have flowered during
the past years.
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The Aeonium populations on Rangitoto is a must see for anyone addicted to succulent flora who is visiting
Rangitoto. I have missed it during my first trip on the island, but have spent a good hour looking at the
plants scattered on a few hundred meters long strip south of Islington Bay wharf, close to the high tide
mark, some less than a meter above it (and I think quite exposed to the salty sea water during storms) on
rough lava blocks at times on almost vertical steeps, until 3 4 meters above sea level. Literally thousands
of plants, most of them apparently being hybrids, on a magnificent display with a surrealistic touch I would
say, most of them confined to the shore by the eastern coastal walkway, but with few plants making it over
the path into the light bush. It takes time to have a thorough look as many different inter-hybridized forms
exist. Rarely two plants look the same. Unfortunately there was no systematic study done on this
population, which thats my 50 cents worth may include unique forms and natural hybrids. I have
collected 5 plantlets, two of them with haworthii, two with undulatum and one possibly with ciliatum
ancestry, but unfortunately have turned rapidly into rather horticultural looking plants after being potted
and placed in my greenhouse. Growing conditions cannot be the same I am afraid.

And here we go again, with the rest of the names:

Bryophyllum delagoense (Ecklon & Zeyher) Schinz [Madagascar] (9) is a plant referred under several
names - Bryophyllum tubiflorum Harvey, Bryophyllum verticillatum (Scott-Elliot) Berger, Kalancho
delagoense Ecklon & Zeyher, Kalancho tubiflora (Harvey) Hamet and also known under a colourful
common name the Lizard Plant. It is a long lasting presence on Rangitoto being collected in 1971 by A.
E. Esler; the name is published only in 1988 by W. R. Sykes in vol. 4 of Flora of New Zealand. It grows on
lava blocks, which is not exactly a match to its original habitat, and is a garden escape from nearby
gardens. There is no other New Zealand known wild population.

Bryophyllum daigremontianum (Hamet & Perrier) Berger 1930 [Madagascar, but also Tropical Asia,
especially Pakistan very probably of cultivated origin] is referred as Kalancho daigremontiana Hamet &
Perrier in Wotherspoon & Wotherspoon (2002) and considered a Class 3 priority plant, with the long term
goal Zero Density to be achieved; odd enough not mentioned by Craig J. Miller (1994) at all.

Bryophyllum pinnatum (Lamaire) Oken [Tropical Asia] () just few isolated plants, but as it is
considered to have an immense invasive potential it has been blacklisted, mentioned to be naturalized by
W. R. Sykes in 1977, but placed for the first time on Rangitoto by D. R. Given in his 1984 Checklist.

Bryophyllum daigremontianum x Bryophyllum delagoense [garden hybrid], also known as Bryophyllum
'Houghton's Hybrid' or Bryophyllum 'Houghtonii' is usually referred by New Zealand authors as Kalancho
daigremontiana x Kalancho tubiflora and apparently very rare (or rather a casual presence I would say).
Mentioned for the first time in Rangitoto in 1995 by C. J. Webb as growing on open slopes and old lava
rubble. No inflorescence of this plant was observed.

Crassula coccinea Linnaeus 1753 [South Africa], also known as Rochea coccinea (Linnaeus) De
Candolle used to be another very popular garden plant few decades ago due to its big bright red flowers
having a tubular corolla, characteristic for the section Kalosanthes. The Rangitoto plants are a classic
garden escape, but very persistent around both wharfs and some of the old abandoned baches. It is not
forming dense population, but rather isolated plants are scattered on the lava blocks. It is definitely one of
the plants enjoying this kind of substrate. It was first collected by W. R. Sykes in September 1982 and is
considered a Class 3 priority plant.

Page 16

Crassula multicava Lemaire 1862 [South Africa], a well known old fashioned garden plant also known as
Fairy Crassula (10), first collected by C. M. Smith in 1957 and mentioned by Healy in 1959, is one of the
most widely dispersed succulent plants on Rangitoto, which is rather sad due to its highly invasive habit. It
grows almost everywhere, on different types of substrates, from bare lava blocks to richer soils in former
gardens, and is the only one being capable to evade the coastal habitats and to penetrate the hostile lava
fields of the interior. I have seen very large mounds here the biggest I have ever seen cultivated or in
nature. It is a Class 2 priority plant, intended to be eradicated on a long term and to be restrained to Zero
Density as a short-term goal, but honestly Im rather disappointed, as the populations seem to increase year
22. Crassula coccinea, a
plant growing close to Yankee
wharf, on the eastern coast of
Rangitoto Island. I saw just a
few plants, not really in their
prime, but this is a winter
growing plant. As 2007 / 2008
was a very dry and hot
summer, the dehydrated
leaves are no surprise.
23. A very large clump of
Crassula multicava
growing close to the entry
of an abandoned bach.
Note the many flowering
stems, developing many
easily detachable plantlets
getting airborne with the
slightest breeze - the main
dispersal vector of this
highly invasive plant.
Page 17
by year. I have seen many flowering sized plants and thats exactly what shouldnt have happened because
of the plantlets formed on the inflorescences and getting airborne to be dispersed over large areas. Huge
mounds of mature plants are an ever-lasting source of tiny plantlets and therefore I am rather skeptical
about the success of the Eradication.

Crassula orbicularis Linnaeus 1753 [South Africa] is a very popular plant here in New Zealand, often
sold in Garden Centers under different names such as Crassula rosularis Haworth or Aeonium x haworthii
and masquerading in collections under these names for decades. It was discovered on Rangitoto just
recently in 2005, few isolated plants persisting in the proximity of the abandoned quarry (which I havent
visited yet) but posing probably a high danger for the habitat. It was blacklisted immediately, I think
because of the very prolific seed production and high germination rate, but also because of the stolons it is
shooting, another very effective way of vegetative reproduction and dispersal. However, action was taken
and all plants were removed and in 2006 no other plants were found when the site was revisited.
Reportedly isolated plants were seen since in few Northland localities, not on Rangitoto though.
Fortunately enough the plant is very sensitive even to light frosts which occur periodically on Rangitoto, so
that it is quite possible that small populations ready to take off to a higher plateau level are decimated back
to few rootstocks or stem fragments. It is unlikely that Crassula orbicularis can survive in nature
unassisted south of Auckland.

Crassula sarmentosa Harvey [South Africa] was known to have escaped in nature since 1989 in
Auckland, but P. J. de Lange has collected it only in 1995 from a slipway in Rangitoto, growing as a
garden discard on scoria blocks. The finding was published in P. B. Heenans 1999 Checklist. Apparently
still very rare, Crassula sarmentosa is not considered a threat and is not included in the current vegetation
control program.

Crassula tetragona Linnaeus 1753 [South Africa] was also first mentioned in 1959 by Healy and having
now a wide dispersal area especially on coastal cliffs near urban areas, but staying in small numbers,
mostly isolated plants. There are just very few places where it forms important populations and one of
them is Rangitoto Island. Odd enough I havent seen the plant in Rangitoto, but in a remote and random
place north of Auckland back in 2003. However, the volcanic fields of Rangitoto are definitely a match for
24. A smaller Crassula
multicava with leaves
displaying clearly stress and
water deficit, but still in
flowering in late January,
long after the typical
flowering period. The nice
flower colour and the
wonderful display of
hundreds of flowering stems
in large plants stand for the
common name of this plant
The Fairy Crassula.
Page 18
this plants needs the preference for this type of substrate you can see in cultivation; put a plant on a heap
of volcanic rocks or in a crack in a stone wall and it will thrive like nowhere else.

Echeveria cv. Set-Oliver Hort. has an uncertain status. There is no botanic collection or formal
identification of plants growing on Rangitoto Island but there is a strong belief that this plant is present
here as a garden escape. As this old garden hybrid is extremely popular here in New Zealand I find it
throughout possible.

Echeveria multicaulis Rose [Mexico] is another plant with uncertain status; it was first mentioned in 1992
as being present only in Rangitoto with only few isolated plants. There is also an A. E. Esler collection
dated 12.09.1971 of a wild growing plant, which is believed to be Echeveria multicaulis although the
herbarium specimen is not numbered and formally identified. C. J. Webb (1995) notes that very few plants
were observed close to an old house in Islington Bay. Unfortunately I have not seen it although I have
browsed thoroughly in this area.

Echeveria secunda W. B. Booth 1838 [Mexico] - small groups are scattered in different New Zealand
localities on both major islands and is reportedly present in significant numbers in Rangitoto, although I
havent seen it. It is a very popular plant here, one of the first Echeveria to be used in landscaping, and as
vegetative reproduction is very easy leafs, stem fragments or loose rosettes are rooting all year round if
the right conditions are given it is actually no wonder that the plant has managed to escape in nature.
There are many herbarium specimens collected in time, but H. H. Allen states that there is some doubt
about the authenticity of all collections, including some of Rangitoto, many of the earlier collections being
confused with Echeveria elegans Rose [Mexico]. Although no Echeveria elegans population has been
documented in New Zealand we cannot discard completely the statement of one of the patriarchs of New
Zealand Botany. Echeveria elegans forms bigger rosettes, has bigger and thicker leaves and is more
glaucous, but these are not characteristics to be retrieved from herbarium specimens.

Echeveria setosa Rose & Purpus 1910 [Mexico] was first mentioned in 1995 as a rare presence on the
island. However, this genus is probably just a colourful and somehow exotic addition to the native flora
and poses no real threat and is not included in any eradication program. A single 1990 collection is known,
made by W. R. Sykes.

Sedum acre Linnaeus 1753 [Europe] was mentioned in 1904 by W. W. Smith and is probably the most
wide spread Sedum in New Zealand. It is present also on Rangitoto reportedly in large numbers, but I have
to admit again that I havent seen it here or anywhere else in New Zealand. The plant is inconsistently
mentioned as present in Rangitoto D. R. Given (1984) and Craig J. Miller (1994) do mention it, but not
Wotherspoon & Wotherspoon (2002). I can only assume it is an omission.

Sedum album Linnaeus 1753 [Europe] has also a wide dispersal area and shows similar habitat needs as
Sedum acre. Collected in 1954 by H. Talbot and mentioned in 1959 by Healy, Sedum album is known to be
present in Rangitoto since D. R. Givens 1984 Checklist. I have seen this plant on large areas in Rangitoto,
apparently dispersing from an old abandoned batch in Islington Bay; literally hundreds of plants growing
among Crassula multicava seem to have been dispersed from what used to be a border strip all over the
path sides southbound on the eastern track, forming from place to place lovely clumps. Most of the plants I
have seen (flowering at the time of my second trip to Rangitoto) were growing directly on the lava blocks
where lichens were already present and very exposed to sunlight, giving therefore a yellow-reddish tint to
the plump succulent leaves. Wothersopoon (2002) is not mentioning the plant again.

Page 19

Sedum forsterianum C. A. Smith 1808 [Europe] is scattered in several localities in Auckland area,
including Hauraki Gulf islands such as Little Barrier Island and Rangitoto. Beever has done the first
collection back in 1978 in Little Barrier Island but the first account of this finding appeared 10 years later,
in vol. 4 of Flora of New Zealand, and in C. J. Webbs 1989 Checklist. It has been confused several
times with Sedum reflexum in New Zealand as some of the distinguishing characters are evident just on
living material and cannot be preserved in herbarium specimens. I havent seen it and it is not mentioned
by Wotherspoon & Wotherspoon (2002) and Craig J. Miller (1994).

25. Sedum album, very
common in Rangitoto,
probably second after
Crassula multicava. It grows
mostly in the proximity of
abandoned baches and
gardens, but it also shows
clearly that it can disperse
successfully some of the
plants I have seen here were
quite far from the obvious
originating sources.
26. Sedum album still
in flower. I wonder
how it looks like
during the typical
flowering period as in
many places the plant
has spread like a
ground cover. Very
nice flowers indeed.
Page 20
Sedum mexicanum Britton 1899 [Mexico] is mentioned by David R. Given (1984) as being present only
on the South Island (Heathcote Christchurch), but this name pops up now very surprisingly in
Wotherspoon & Wotherspoon (2002) as a Class 3 priority plant targeted to be eradicated from Rangitoto. I
havent seen it and I also couldnt find any reference in botanical literature or in other plant control
programs placing this plant in Rangitoto. However, in case it wasnt confused for Sedum reflexum, I think
it is very likely to occur here, as Sedum species are particularly well suited for Rangitoto habitats, and very
popular in cultivation as well.

Sedum praealtum ssp. praealtum De Candolle 1847 [Mexico] is another Mexican Sedum originating from
alpine habitats, which is very widespread in New Zealand. It was first collected in 1953 near Whakarongo -
Bunnythorpe (Manawatu) but in short time other populations were discovered so that in 1959 Healy
already knew of its extensive colonies formed in both major islands. It is quite common on Rangitoto
Island, where I have seen more or less isolated plants scattered all the way from Islington Bay wharf
southbound on the coastal track until it leaves the shore to cross the bush land to Rangitoto wharf.

27. My younger son Vlad
with one of the Sedum
praealtum ssp. praealtum
he has seen on the lava
blocks close to Yankee
wharf. This is one of the
older plants; most of the
plants we have seen were
young plantlets 1 2
years old.
28. Two Sedum praealtum
ssp. praealtum plantlets
growing between debris and
decayed lichens. Most of the
plants I have seen here were
of this size. It is said to be
quite common, but I have
seen more or less isolated
plants and no significant
populations.
Page 21
Sedum reflexum Linnaeus 1753 [Europe] was first mentioned by Healy in 1959 but was already known
from a 1954 Otago collection. Apparently it has spread since, and in D. R. Given (1984) records the second
distribution area in Gulf Hauraki (Auckland) on several islands including Rangitoto Island and Little
Barrier Island. It is another abundantly established Sedum growing on rocky outcrops and coastal cliffs. It
is mentioned by Craig J. Miller (1994) but not by Wotherspoon & Wotherspoon (2002) but I didnt see it.

In the end Rangitoto Island is a fascinating new world worth more than a trip or two, its a piece of
natural history unveiled to our eyes, and definitely one of the most exciting places here in New Zealand for
succulent freaks like me. With 25 28 succulent vascular plants present here (11), some fully naturalized,
some possibly just casual, Rangitoto Island is definitely a paradise for succulent flora, even for endemic
species such as Disphyma australe ssp. australe, which is present on the island.

29. We left Rangitoto Island on a dull
rainy evening with a tropical storm
threatening to batter Auckland during
the night but still we could call it a day!
Page 22
Additional References:

David R. Given Checklist of Dicotyledons Naturalized in New Zealand Crassulaceae, Escalloniaceae, Philadelphaceae, Grossulariaceae,
Limnanthaceae (New Zealand Journal of Botany, Vol. 22, 1984);

P. B. Heenan & al. Checklist of Dicotyledons and Pteridophytes Naturalized or Casual in New Zealand: Additional Records 1994 1996 (New Zealand
Journal of Botany, Vol. 36, 1998);

P. B. Heenan & al. Checklist of Dicotyledons, Gymnosperms and Pteridophytes Naturalized or Casual in New Zealand: Additional Records 1997 1998
(New Zealand Journal of Botany, Vol. 37, 1999);

P. B. Heenan & al. Checklist of Dicotyledons, Gymnosperms and Pteridophytes Naturalized or Casual in New Zealand: Additional Records 1999 2000
(New Zealand Journal of Botany, Vol. 40, 2002);

Craig J. Miller & al. ARK2020: a Conservation Vision for Rangitoto and Motutapu Islands (Journal of The Royal Society of New Zealand, vol. 24,
1994);

S. M. Timmins & H. Braithwaite - Early Detection of Invasive Weeds on Islands (2002 or more recent);

C. J. Webb & al. Checklist of Dicotyledons, Gymnosperms and Pteridophytes Naturalized in New Zealand: Additional Records and Corrections (New
Zealand Journal of Botany, Vol. 27, 1989);

C. J. Webb & W. R. Sykes - Checklist of Dicotyledons, Gymnosperms and Pteridophytes Naturalized or Casual in New Zealand: Additional Records 1988
1993 (New Zealand Journal of Botany, Vol. 33, 1995);

S.H. Wotherspoon & J.A. Wotherspoon The Evolution and Execution of a Plan for Invasive Weed Eradication and Control, Rangitoto Island, Hauraki
Gulf, New Zealand (ca. 2002);

Further Readings:

Jacqueline Crompton Ottaway - Rangitoto - Auckland's Fragile Icon (2004);

A. E. Esler Botanical Features of Motutapu, Motuihe, and Motukorea, Hauraki Gulf, New Zealand (New Zealand Journal of Botany, vol. 18, 1980);

Alastair Jamieson Rangitoto (New Zealand Geographic, 2004);

My Notes:

(1) Photographs from the second half of the 19
th
century are showing very clearly that the vegetation on the island was far much reduced compared to what
it is nowadays and probably far less diversified. Bare lava fields were extensive throughout the island almost up to the top of the volcanic cone. Even today
the vegetation is not that rich and dense being unable to support large bird populations. Although the crucial importance of this natural sanctuary has been
widely acknowledged in the first half of the 20
th
century, structured and sustained researches and conservation programs have started only with the late 50-
ties early 60-ties.

(2) Zero Density means disposing of the plants before reaching the adult stage and before sexual reproduction becomes possible. Only vegetative
reproduction is possible (which in these cases is less efficient) posing no real threat for the habitats.

(3) Refer to Part 1 for a regional map of New Zealand.

(4) The description of the entire process will be part of another article, entirely dedicated to Rangitoto Island.

(5) The climate is mild and essentially different from the adjacent areas, even if the rainfall levels seem to be marginally higher than the 1,268 mm / year
recorded at the nearest meteorological station Albert Park in Auckland City. Some light frosts may occur during the winter but the average temperatures
are ca. 11 degrees Celsius in July and 20 degrees Celsius in February; the number of sun hours exceeds 2,100 per year (49% of the possible number) and
strong winds are not very often.

(6) I was mentioning this number on few occasions in previous articles, and this appears to be an error. However, there is always a discrepancy between
numbers raised by environmental entities (and the Wotherspoon and Wotherspoon paper is a report for the Department of Conservation) which are not
always 100% substantiated and the botanical accounts based strictly on confirmed botanical collections, but which may lack on field study efficiency or on
local, regional and national adequate cover. Fact is that I have seen and documented on Rangitoto Island at least two species, namely Agave americana and
Aeonium haworthii, not mentioned in botanical records, but included in eradication and control programs supervised by Auckland Regional Council and /
or the Department of Conservation.

Page 23
(7) I have seen in 2002 or 2003 a very impressive variegated specimen on a beach just north of Mt. Maunganui, on the sand dunes concealing the lagoon.

(8) Aloe latifolia is considered nowadays just a minor variation from the type plant having broader leaves. The name fell into synonymy many years back.

(9) No matter how one or another of the botanists or taxonomists may consider I have used in this article for practical reasons the name Bryophyllum and
not Kalancho as this name seem to be more often referred by New Zealand authors.

(10) You can read more about this plant in Part 1 of this series.

(11) I am pretty sure this is not an exhaustive list.

------------------------------------------------

All errors, omissions and misconceptions are mine.

All photos by Eduart Zimer, except 9 & 10 by Vlad Zimer.

Eduart Zimer, July - August 2008

http://eduart.page.tl/Home.htm


Eduart Zimer - (SDU) - Adventive Plants - Part 4 (2008)

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Succulent plants from down under Adventive Plants (Part 4):

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