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Succulent plants from down under - Disphyma australe and its natural hybrids Although the very most

of the Aizoaceae are growing in South Africa and Namibia (actually one cannot think one without the other) there are few genera belonging to the group called by the informal name of mesembryanthemums which are naturally occurring in several regions of the Southern Hemisphere Carpobrotus N. E. Brown seem to be the prototype growing from the coastal regions of western South Africa to Australia and Tasmania and to the Americas as well (the coastal regions of California and Chile), but also Sarcozona J. M. Black and Disphyma N. E. Brown are well represented in Australia and the Southern Pacific. Even the geographical distribution of these plants is pointing towards a very likely gondwanic origin of the Aizoaceae, but this is another story. One of the plants belonging to this group Disphyma australe (Aiton) J. M. Black, a New Zealand endemic, is a very interesting plant to say the least. There was (and still is) a bit of confusion around this name, as some botanists consider that more study on the genus Disphyma is still needed, and in order to know exactly what are we speaking about we have to do if we like it or not, but mostly not due to the ever and ever changing names of the plants we are interested in some taxonomic research. From the actual view on the species outlined by R. J. Chinnock, Disphyma australe (Aiton) J. M. Black 1932 sensu lato (1) is grouping quite different plants, some growing in Australia, some in New Zealand or even on remote islands of the Southern Pacific, not to speak of parts of this complex being merged with South African relatives. Due to the intensive work and field research of R. J. Chinnock (1971, 1972, 1976) we have today a different view. Based on the study of the capsule structures and other vegetative and floral characters Chinnock has segregated the historic Disphyma australe complex in three distinct species: Disphyma clavellatum (Haworth) Chinnock 1976 this name refers now the Australian populations. The synonyms to this name are: Mesembryanthemum clavellatum Haworth 1803 ; Disphyma australe (Aiton) J. M. Black 1932 sensu lato; Disphyma blackii Chinnock 1971 and in more recent years Disphyma crassifolium ssp. clavellatum (Haworth) Chinnock 1986. As Chinnock recombined back in 1971 Disphyma blackii separating the Australian populations due to very specific differences from the rest, he omitted the fact that a name was already available for the plant he was describing. This name, never referred before by any Australian author says Chinnock, is Mesembryanthemum clavellatum. Haworths description was based on plants raised from seed collected by Robert Brown in Australia apparently from cultivated plants, but anyway he is stating in his unpublished diary that the original specimens were collected at Goose Bay in southwest Western Australia. Although no Goose Bay population is actually known, Chinnock considers Haworths account of the species quite sufficient to distinguish the plant from other Australian mesembryanthemums. Five years later Chinnock makes the correction and recombines the taxon using the historic attribute. Im not insisting on this, but its worth mentioning that in more recent years it was considered a synonym of Disphyma crassifolium ssp. clavellatum in other words it is now considered a subspecies of the South African Disphyma crassifolium. To add to the confusion Dr. H. E. K. Hartmann (2000) considers firmly that there are two different species, Disphyma crassifolium and Disphyma clavellatum; but Chinnock, the author of the last named taxon takes now a step back and considers that the Australian populations are not that different to warrant a complete separation from the South African species. As Didge Rowe puts it this needs more study (2). Disphyma papillatum Chinnock 1971 this name refers now the Chatham Islands populations. Although restricted to a rather small area this plant has a greater variability than Disphyma australe, especially regarding the leaf and flower pigmentation. There are not only green forms
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or red forms but some of the red ones tend also to have a more or less distinctive brownish touch. Flower colour is also very variable ranging from white to pink and to purple with all sort of intermediate nuances. As colour variants are not restricted to a certain population but all sort of forms may occur within a single population it makes no sense to divide further but to accept its natural variability. A characteristic of this species unlike the other plants mentioned here, it has always acute leave tips. Disphyma australe (Aiton) J. M. Black 1932 sensu Chinnock this is the plant we are interested in. During his research on Disphyma in the late sixties and early seventies Chinnock has observed several variations within different individual plants and populations regarding the pigmentation but also, more important, the leaf margins of some populations from the Kermadec group of islands (Raoul, Curtis, Macauley and Lsperance). Based mainly on this variation Chinnock (1976) splits the taxon into two different subspecies: o Disphyma australe (Aiton) J. M. Black 1932 ssp. australe endemic to New Zealand and scattered almost all over the coastal zones from the north of the Northern Island to the southern parts of the Southern Island (except some of the very southern extremities and the Stewart Island, probably not because of the cooler climate as we will see, but because of the quite different habitat) with just few major interruptions mainly south of Greymouth on the western coast of the South Island, but also on the eastern coasts. It occurs also on Chatham Islands (H. H. Allen, 2004). Its only synonym is Mesembryanthemum australe Aiton 1789. o Disphyma australe (Aiton) J.M. Black ssp. stricticaule Chinnock 1976 endemic to the Kermadec group of islands, some 1,200 km north-east of New Zealand, very isolated in the Southern Pacific. In recent years this subspecies was introduced (possibly by man) in New Zealand and it occurs now in few locations growing wild together with the type plant, but it needs more warmer conditions. Its not that common in New Zealand, rather rare I would say, but quite abundant in some of the Kermadec group of islands (it is the dominant species on Curtis Island, but less common on Raoul Island). A very interesting feature of the cultivated plants of this subspecies the new stems have always an erected growing pattern for some time but becoming prostrate and spreading later on; this growth pattern was not observed at all in wild populations. This plant seems to be the most isolated Aizoaceae. In fact there is a quite strange difference (or incompatibility I would say) between the two subspecies, as pointed out by Chinnock, quite amazing considering that both subspecies are vegetative extremely similar, but I will keep this for later on. Disphyma australe is a spreading and mat forming plant with long and thin stems, reaching usually to 50 mm in length, but sometimes branching and growing to 100 cm long or even more and usually to 5 mm in diameter, sometimes even more; the stems are usually green but younger parts of the stems may have a reddish colour. The stems are forming from place to place some nods from where the succulent leaves are growing; from the same nods the adventives roots may appear. The internodes are varying a lot usually from 0.5 to 5.0 cm in length, but reaching sometimes even 10.0 cm. Axilary short shoots are condensed and terminating usually in flower but may also develop in long stems. The leaves are succulent and elongated, and connate towards the base and somehow with angled edges, but with smooth margins. The leaves are growing in slightly unequal opposite pairs or more often in opposite clusters and are usually 20 50 mm long, sometimes even longer and about 6 mm in diameter,
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occasionally up to 9 mm, and have the tips pointed. Usually the sides of the leaves orientated towards the stems are somehow smoother. The flowers can have 20 40 mm diameter and can vary quite largely from white to pink. The flowering period in habitat is from late winter to early summer and open in late morning for several days and closing during the night; the flowers are opening for several days in a row and with each day the colour is getting somehow darker, even the whitish ones. Experienced growers can say for how many days the flower is open. Occasionally few flowers can be borne till early autumn. The fruit consists of a dry capsule, usually 5 8 celled, containing numerous small brown to dark brown seeds (0.9 1.1 x 0.6 0.8 mm). Several small variations were noticed (mainly flower colour, stem colour, but also leaves form and colour), but not enough marked to recognize them taxonomically as distinct forms. Sometimes variations are occurring only due to different environmental conditions. Chinnock (1971) states that extreme environmental conditions are leading to variations in leaf length, pedicel length, flower and capsule diameter, internode length and pigmentation. The most usual deviation from the normal is that seen in shady or wet places where leaves and pedicels are long. High exposure to sunlight and wind, or shallow soils causes reductions in size of the various parts. Very reduced plants are found on open and exposed rocky platforms. However, there are two different variants of the plant, one green (never producing red colour under any circumstances) and one, which contains the red pigment (betanin) in different amounts. Based purely on the flower colour there might be considered two or three different forms, partly having a distinct distribution, but also growing together in few locations (3). The red variant of the plant is always bearing light or dark pink flowers and the green one always the white flowers. The flower is selfsterile, so that cross-pollination is needed in order to set seed.

1. Disphyma australe ssp. australe (Muriwai, February 2007) (Photo: Eduart Zimer)
This is a photo of a plant taken in February 2007 in Muriwai (quite close to Auckland) on the eastern coast (The Tasman Sea). As you can see it is a well grown plant, hidden between grasses and other plants in a quite sheltered place on the plateau, at some 15 to 25 m above the sea. It has definitely longer leaves as those in Nicks photos, must be some 5 cm long or even longer. The place is quite close to the spot from where I have done the photo in which you can see the sea (photo 2 Muriwai habitat). There were just few small patches of plants around, none bearing flowers unfortunately. Such small groups are scattered everywhere along the coast here, I havent seen big populations.

Chinnock maintains the historic attribute for the New Zealand populations. Mesembryanthemum australe was first collected by Joseph Banks and Daniel Solander, and illustrated in 1769, during
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Cooks first voyage and mapping of New Zealand coastlines. Later, in his unpublished manuscript (hence sometime the first description is erroneously attributed to Solander) Primitiae Florae Novae Zealandiae dated 1773; Solander states that the plant is abundant close to the sea especially in rock fissures near Tolaga, Opuragi, Motuaro, Totaranui, New Zealand. Aiton published the name in 1789 and stated that it was a New Zealand native plant. He possibly has used for his diagnose a cultivated plant from Kew as it is known from his own account that it was already introduced to Britain since 1773. J. M. Black recombined in 1932 the taxon including it in the quite recent created genus Disphyma N. E. Brown 1925. Disphyma australe was originally considered by J. M. Black to occur in Australia, Tasmania, New Zealand and some other South Pacific islands. In early botanical accounts Mesembryanthemum australe is also recorded on Lord Howe Island and Norfolk Islands. Chinnock has not found in his field trips any evidence, nor has listed Turner the plant in his 1968 Norfolk Islands plant list. On the contrary, Chinnock believes that the plant was originally confused with a smaller form of Carpobrotus glaucescens, although the later one bears a very distinctive berry-like fruit, so that all these accounts should be discarded. Even if restricted to the coastal line Disphyma australe is quite common and sometimes even abundant in many dozens of locations scattered all over New Zealand, sometimes hanging on rock cliffs of the mainland or off-shore islands or growing beside grasses (usually associated with Poa astonii or Poa anceps) on the rocky soils or gravels of the nearby plateaus, from sea level to 50 m altitude or more on occasion. Even if it is not a typical sand dune plant it grows occasionally in such places. It can grow successfully basically everywhere near the coast but the salt marshes. It not only survives a very strong marine exposure (not typical for the usual succulent plants, not to speak of the Aizoaceae) but it simply loves it it does not grow far from the shores and is quite hard to keep alive and well in the more arid and hot conditions of the inland.

2. Muriwai habitat, February 2007 (Photo Eduart


Zimer)

This is a typical coastal zone, with steep cliffs and plateaus. The plants photographed by me were somewhere in the area behind me, all growing together with grasses or small vegetation, more sheltered than Nicks plants as you can see below

In his account on Wharekakahu Island vegetation T. R. Partridge (1983) makes an excellent description of a typical habitat of the plant and the usually associated plants: The island is surrounded by a zone of seaweeds typical of the exposed rocky shores of Otago. () Rock pools occur only in the northwest corner of the island where there is a protected rock platform. These pools contain the conspicuously
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green Enteromorpha sp. Cliffs rise about 40 m to a gently sloping plateau. In many places these cliffs are too smooth and steep for plants, but elsewhere there are ledges supporting tussocks of Poa astonii, with occasional Disphyma australe and in moist crevices, Crassula moscata. As the slope angle decreases, the soil becomes deeper and the cliff vegetation grades into that of the plateau. The central third of the plateau is bare rock with pockets of soil in small hollows. () The only common vascular species in the small pockets of soil is the prostrate Atripex buchananii. The rocks are encrusted with lichens. On the slopes of the eastern and western sections there is deep mineral soil, which is modified by many wide-mouthed burrows of nesting sea birds. The vegetation is grassland of Poa astonii tussocks with Disphyma australe. Other frequent plants are Lepidium oleraceum and around the base of the tussocks, Lolium perenne. () On the flat part of the western third of the island bird burrows are present but considerably fewer than in the grassland. The vegetation is dominated by the shrub Hebe elliptica (). Where gaps occur () tussocks of Poa astonii are still plentiful, but beneath the canopy of the shrubs the ground is bare. Vegetation types which are almost the same have been described by Fineran (1966) on Bird Island and Johnson (1976) on Womens Island. In both instances Poa astonii, Disphyma australe, and Lepidium oleraceum vegetation is plentiful on the sloping cliffs with Crassula moscata in crevices and Hebe elliptica on exposed but flatter plateaus.

3. Disphyma australe ssp. australe (Palliser Bay Wellington, 1995) (Photo Nick Perrin)
This photo appeared Nick Perrins 1996 article. It grows directly on sediment rock (a degraded sandy silt) and as you can see has a more compact growth, with shorter leaves. All Palliser Bay photos were taken in October. It is a pinkflowering form.

In several locations Disphyma australe is quite abundant. Sometimes it is even the dominant form of vegetation, as it is on White Island, where it dominates the herb fields and grassland together with Einadia trigonos ssp. trigonos and Poa anceps ssp. anceps all of them primary succession vegetation types (4). White Island is still volcanic active - this means not necessary lava eruptions or lava flow but fumes or solfataric activity with small steam and tephra eruptions from time to time. It is a 238 ha andesite cone which had between 1976 and 1981 the largest and most intense eruptive activity recorded in recent times, destroying partly the vegetation on the island. This is a very good example to illustrate how opportunistic this plant can be, replacing a well settled population of dominant vegetation. A survey conducted by Bruce D. Clarkson (1990) on this island after the major volcanic activities has revealed that Disphyma australe is able to recover and occupy any free niche (due to volcanic activity or even land slides) replacing the former dominant species in a very short time (10 to 20 years) even if
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a seed bank from the previous years is present in the soil for the other species as well. Some other vascular plant species were probably highly disturbed by the increased volcanic activity in the later years and may have simply disappeared from the island. The vascular flora on White Island was represented between 1915 and 1967 by ca. 13 different species, but Clarkson couldnt record more than 7. There are also several other factors allowing the plant to spread quickly when the opportunity is there for example very strong winds bringing seawater into the air and misting the entire habitat. This phenomenon may occur now and then or every few years, depending on the region. Under these circumstances Disphyma australe can replace Hebe elliptica, a companion plant in most of the habitats and very sensitive to seawater, as a dominant plant. This phenomenon may act as a double edged sword as strong winds and heavy rainfalls may spread around in the area the guano from the sea birds usually nesting in the habitat and under these circumstances entire patches of well settled plants of Disphyma australe may be wiped out in a matter of days (although contradictory information regarding the tolerance to guano exists). On the other hand Disphyma australe is very exposed to mechanic disturbances as goats grassing for example entire populations can be destroyed on a grassed land within few years. It is also very sensitive to trampling, once disturbed and damaged the plant is ceasing the rapid spreading for some time.

4. Disphyma australe ssp. australe (Palliser Bay Wellington, 1995) (Photo Nick
Perrin)

The photo appeared also in Nick Perrins 1996 article. It grows directly on the same sediment rock as the plant above and as you can see has a more compact growth, with shorter leaves, possibly more exposed to winds and harsh sun. It is a pink-flowering form. Shows very good the habit of the plant.

This plant has been naturalized in few other places such as the western coast of Australia where is found growing in places with strong oceanic influences. It is rather rare though. It also grows in nature in few of the South Pacific islands especially in Chatham Islands. The climatic conditions are important in order to understanding this plant. Few climate statistics first: North Island: Waitakere (close to Muriwai Beach, but more in inland) July 9 / 14 degrees Celsius, February 17 / 24 degrees Celsius, rainfall of 898 mm / year, almost evenly through the year but just a bit more abundant in May July); Whangarei Heads July 10 / 15 degrees Celsius, February 17 / 23 degrees Celsius, rainfall of 1,362 mm / year almost evenly through the year but just a bit more abundant in December January and June August; Mt. Maunganui July 7 / 14 degrees Celsius, February 16 / 24 degrees Celsius, rainfall of 1,093 mm / year, more abundant in April July; Wellington Harbour July 8 / 12 degrees Celsius, February 15 / 21 degrees Celsius, rainfall of 748 mm / year, evenly through the year.
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South Island: Tahunanui July 2 / 12 degrees Celsius, February 13 / 22 degrees Celsius; rainfall of 920 mm / an almost evenly through the year, but surprisingly June and July (the winter months here) are rather dry months; Dunedin (city placed in the middle of the Otago coastline) July 3 / 9 degrees Celsius, January and February 10 / 19 degrees Celsius, rainfall of 937 mm / year, evenly through the year; T. R. Partridge states that in Wharekakahu the rainfall is of 758 mm / year; again T. R. Partridge at Taiaroa Head 723 mm / year and a mean annual temperature of 10.6 degrees Celsius; J. B. Wilson at Nugget Point rainfall of 852 mm / year and at Tautuku further south 1,214 mm / year, in both cases evenly spread through the year. First of all the rainfall appears to be in most cases if not abundant (occasionally even up to 2,300 mm / year) then at least sufficient for a constant water supply, but it isnt quite so considering that Disphyma australe is growing mostly on rocky superficial soils, very permeable in most of the cases, or even cramped directly on steep rocky cliffs, with poor water retention if any. Also the harsh sun and the strong winds are drying out the soil quickly so that even after storms and heavy rainfall in a matter of days this may become bone dry at least on the surface. There are in some places (or in some years) even short periods of draught. There are also big differences in the number of sunny days in a year between different locations, not to speak of the number of rainy days varying greatly from 150 to 250 / year. A high number of rainy days (with more than 0.2 mm recorded rainfall) does not necessary mean a high level of rainfall but indicates a constant water supply and may also point towards moderate light conditions (bright light but not too sunny) as being acceptable for the plant, in contrast with the Whangarei Heads for example one of the sunniest places in New Zealand.

5. Disphyma australe ssp. australe (Point Jerningham Wellington, ca. 1996) (Photo Nick
Perrin)

Again a Nick Perrin photograph taken probably in 1996. Very nice long reddish stems hanging down from the cliff.

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It is interesting to see that Disphyma australe does thrive in quite different temperature conditions, unless its not too hot or its not freezing; it grows happily from the far north of the Northern Island (where annual mean temperatures are reaching 16 degrees Celsius and frost is inexistent) to the southern parts of the South Island, where annual mean temperatures may drop under 10 degrees Celsius and light frosts may occur even on the coastline. Although it is said that the plant can resist for short time at exposure to -4 degrees Celsius, relying strictly on generally available climate statistics may not show exactly the truth. In their account of the Catlin cliff vegetation J. Bastow Wilson and Carol Cullen (1986) are showing that in this region of Otago, Southern Island, readings of the Nugget Point station (placed on the coast) are showing a mean annual temperature of only 9.9 degrees Celsius. At Tautuku and Owaha further south, where the stations are in the inland there are 13 and 43 respectively frosty nights per year, but in the proximity of the coast only 3. Minimum temperatures may reach -3.0 and -6.1 degrees Celsius respectively at the two inland stations but only -0.8 degrees Celsius at Nugget Point. Even 10 km towards the inland can make a huge difference. That shows that the plant even if it is not exposed to extremes on the coastline habitats enjoys actually a wide range of conditions, this span of at least 6 degrees Celsius for the mean annual temperatures being a lot! The only thing in common is the marine exposure.

6. Disphyma australe ssp. australe (Palliser Bay Wellington, 1995) (Photo Nick Perrin)
This is a very interesting photo showing both colour variants together, and again this photo has been presented in Nick Perrins 1996 article.

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Cultivation can be quite tricky if not the right conditions are given. It is amazing how limited by the coastal conditions this plant is. Under the right circumstances (read in the proximity of the shore) the cultivation is extremely easy, the nature does it all by itself, of course if its grown outdoors. It becomes more like a challenge to grow it in more inland like conditions. Didge Rowe (2006 / 2007), a connoisseur of the Pacific Aizoaceae admits that he had problems in keeping his plants in a good shape due to the summer heat and the extremely dry air in Australia (5). It is actually not a good plant for greenhouses as it doesnt like especially the dry air and the high temperatures developed indoors on occasion, but more for an outdoor rock garden. First of all the plant needs a good light; no amount of bright light is too small, even with 4 6 hours of direct sunlight daily although it prefers a slightly filtered or diffuse light. It doesnt grow at all in shadow and it also loves high UV levels. As a shore plant exposed to breezes and sometimes to strong winds it also needs constant ventilation, this cannot be replaced by anything else. Disphyma australe thrives in a loose and almost mineral soil, very light and with very little organic components and although it is not too hard to meet in cultivation the proper pH needs as it grows happily in slightly acid to neutral to slightly alkaline soils (6) ; it does not like organic soil at all. Under these circumstances it produces a very poor growth and is very prone to pests and diseases and it wont survive even the slightest frost. It also demands a perfect drainage, once waterlogged the plant can die within weeks. Heaps of coarse mineral stone chips and volcanic scoria or pumice, some coarse pumice sand and a bit of loam and just a touch of humus with small pine bark chips will do a good soil. A well grown plant may reportedly resist up to -4 degrees Celsius for a short time, but I wouldnt recommend such an exposure. It also does not stand too high daytime temperatures in summer, 25 degrees Celsius is the higher limit, and over this the plant ceases growing. Exposed regularly to over 30 degrees Celsius Disphyma australe may scorch or even die in short time, even if all other right cultivation conditions are present. It simply cannot stand prolonged heat. Some growers recommend from time to time a slight misting with seawater, which makes sense as in its natural habitat the plant is exposed to strong winds bringing sometimes water from the sea (7). Misting is anyway a good idea especially during the summer months, as the plant loves humid air and rather dry soil. Easy with the water, but do it regularly as the plant does better with a constant water supply. Nick Perrin (in a personal message) states: I have found D. australe difficult to cultivate. It needs very deep pots with permanent dampness at the bottom, I think. In habitat it is often on rocks with roots going about 2-3 m through cracks in the rock down to permanent dampness. It does not seem to like shallow pots that dry out. Disphyma australe can be cultivated as an annual plant even in places where it is not hardy during the winter because it is flowering and setting seed within a year. Producing new plants from seed is easy germination takes place at 8 15 degrees Celsius and the seedlings are growing very fast, but it must be sown very early in the spring as soon as the frosty nights are over. Another method is to take clean cuttings (10 20 cm from the tips of the branches, having at least two nods), keep them in a dry and cool place over the winter and place them in early spring on the soil; as soon as they get some moisture they will root and start grow quickly. The flowering period in cultivation may differ slightly from the period it usually flowers in habitat, in some of the cases when it is grown as an annual plant it may flower in late summer or even early fall. It is not exactly a pot plant, but more one for a raised bed or a rock garden. The plant, also known under the common names of Horokaka or Maori Ice Plant, has edible leaves and has been used for centuries in the traditional Maori culture as a reliable food supply but also as a traditional medicine as it was used to heal burns and superficial skin wounds. Although not that powerful it was used probably in a similar way to the renowned Aloe vera. Very little is known though. The meaning of the name Horokaka is not exactly known. Horokaka is also the name of ceremonial rites of war involving fire, but a meaning of the word horo is landslide, and with kaka meaning
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hair would make more sense considering that the plant is occupying very easy bare ground disturbed by landslides. However, Horokaka is a very popular plant here down under; a symbol of the New Zealand succulent flora and it is featured on Cactus & Succulent Society of New Zealand badge.

7. The Cactus and Succulent Society of New Zealand badge featuring Horokaka
This is the official badge of Cactus & Succulent Society of New Zealand (CSSNZ) featuring Disphyma australe. Off the records, it pictures actually a non-existent plant I think a pink flowering form with complete green stems and leaves (but dont tell anyone, this will be our secret).

There are two known natural hybrids of Disphyma australe ssp. australe in New Zealand, but there is something very strange here. Although Disphyma australe ssp. australe and Disphyma australe ssp. stricticaule are growing together in few places sharing the habitat they do not seem to hybridize, not even the two distinctive forms of Disphyma australe ssp. australe seem to interact in places where the two are seen growing together, quite strange considering how small the differences (at least the vegetative ones) between the two subspecies really are. In fact it does not hybridize with any of the Disphyma species, even though as a naturalized plant in some coastal regions of Australia grows together with other Australian native Disphyma species. At some point R. J. Chinnock was contemplating even the idea of splitting Disphyma australe from the rest of the genus and creating a new genus for what he was considering to be the only evolutionary development of a new genus in Aizoaceae outside the south-african core of this family, because of the sufficient morphologic differences (flowers, fruit) from the diagnose of genus Disphyma as outlined by N. E. Brown in 1925. Weird enough Disphyma australe ssp. australe is hybridizing very easily with two remote relatives Carpobrotus aequilaterus (Haworth) N. E. Brown 1928 (synonym Carpobrotus chilensis (Molina) N. E. Brown (8)), growing as the later name states in South America on the Chilean coastline and Carpobrotus edulis (Linnaeus) L. Bolus 1927, a well known south-african relative, which rises a still unanswered question why? So far I know no real answer was given yet. Didge Rowe tries to explain this curiosity due to the existence of a recessive gene, but without any scientific research supporting this, this is just a speculation. Of course it might be very true, but what if instead of being an evolutionary development from the genus Disphyma our New Zealand endemic is just the link between Disphyma and Carpobrotus? Probably in near future phylogenetic studies will tell us the truth. Carpobrotus aequilaterus and Carpobrotus edulis were introduced in New Zealand by man some 150 160 years ago as garden plants 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. Carpobrotus edulis is quite common in many places and without seeking for the plant I have noticed its presence in places as Muriwai, the sand dunes of Ninety Mile Beach or even on sandy patches of very popular beaches such as Long Bay. Unfortunately at Ninety Mile Beach I have seen the plant outside the flowering period, at least on the endless sand dunes of where it grows here in abundance it is for sure a spectacular show in late spring.
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8. Carpobrotus aequilaterus, unknown location 2002 (Photo Nick Perrin)


I have no idea about the location or time of the year. Nicks elbow ruins the photo but I have to include it here as it is the only photo I have with a confirmed Carpobrotus aequilaterus.

Carpobrotus edulis is a South African Aizoaceae originating from the coastal lines of the Cape Regions. It is a mat forming trailing plant with stems as long as 600 cm, glabrous and 2-angled and sub-woody at the base. The succulent leaves are smooth and sharply 3-angled, 70 120 mm long and 12 17 mm in diameter, keel denticulate at least near the apex. The flowers are quite big, reaching to 80 100 mm diameter when fully open, with numerous spreading 30 40 mm long petals and usually pale yellow but turning pinkish orange with age (9). The longest sepals are 40 55 mm. The small brown obovoid seeds are 1.0 1.3 mm long. The plant grows in almost all coastal regions of New Zealand North Island: Northland, Auckland, Bay of Plenty, Wellington, Wairarapa; South Island: Nelson, Marlborough, Canterbury, Otago, Southland, on cliffs and sand dunes (it is a typical sand dune plant) but it found its way to more inland sites along the railway and roadside cuttings. This show clearly how opportunistic this plants is as it finds excellent niche opportunities in places where the settled vegetation is disturbed by the earthworks. However, it is more adapted to inland conditions and is not depending strictly on marine exposure. It is even grown successfully in Alfriston Botanic Gardens, some 20 km from the east coast (the influence of the shallow waters of the concealed Manukau Harbour at north-west does not really count), a distance that makes quite a difference. The plant is very common in New Zealand, used sometimes in the past on purpose mainly to fix the sand dunes an ongoing ecologic problem here, as it has a more rapid growth and spread than any other endemic plant (10).
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9. Carpobrotus edulis (Muriwai, February 2007) (Photo Eduart Zimer)


This time it is my elbow visible in the right corner. The only flower Ive seen at that time of the year. Usually flowers are bigger, like a tea cup or so; this end-of-the-season flower wasnt even fully open. This flower had 5 cm or so diameter, possibly reaching 7 - 8 cm fully open.

The natural hybrid Disphyma australe ssp. australe x Carpobrotus edulis although scattered almost all over New Zealand coastal lines, present in many locations following quite close the mixed populations of its parents (North Island: Northland, Bay of Plenty, Manawatu, Wellington; South Island: Marlborough, Canterbury, Otago, Southland), was first mentioned by A. J. Healy (1959) under the name of Mesembryanthemum sp. at various coastal locations in Otago, South Island. Healy stated that the identity of the plant couldnt be determined precisely but he assumed that this plant would be closely to Carpobrotus chilensis. It is again R. J. Chinnock to be credited for the correct identification of this plant, due to his 1968 fieldwork. Chinnock suspected the hybrid nature of this mystery plant after realizing that there is no single location where this plant grows alone, but only in overlapping populations of both parents or at least with one of them. In his work on these hybrids R. J. Chinnock (1972) presents an excellent picture of both parents and the hybrid plant. It is worth to see this picture and to download his excellent article available on the Internet. The plant shows very clearly intermediate characteristics the stems, usually 100 200 cm long are weakly 2-angled, the leaves are
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succulent and 20 60 mm long, keel weakly denticulate at apex or entire. The sepals are 15 21 mm long. The flowers have 45 60 mm diameter, the petals are either orange-pink or yellow to whitish, pink with yellow base on ageing. The pollen is mostly highly sterile and fruits are not formed (11).

10. Disphyma australe ssp. australe x Carpobrorus edulis (Muriwai, April 2006) (Photo Eduart Zimer)
A hybrid plant in more or less the same spot as photo No. 1 (not too far anyway) in April. No flowers in sight. The photo shows quite clearly the intermediate form of the hybrid. It is one of the 2 or 3 hybrid plants I have seen; in 2007 I couldnt find it again.

The natural hybrid Disphyma australe ssp. australe x Carpobrotus aequilaterus is found in just couple of places (Patea, Castlepoint) and was mentioned for the first time in 1934 by Cockayne and Allan who were suggesting that certain intermediate forms found near Castlepoint may be due to hybridism with the naturalized Carpobrotus aequilaterus. It is similar to Carpobrotus edulis Disphyma australe but differs mainly because of the flowers having only 40 - 52 mm diameter; the sepals are 12-21 mm long; the petals are purplish, later becoming purplish pink. This compatibility of Carpobrotus with Disphyma is rather strange, as an extreme morphological difference exists, but also the strange ability to hybridize. Chinnock (1972) states: In recent years the classification of the mesembryanthemums has been based upon the structure and form of the fruit. The
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majority of genera are characterized by dehiscent (hygrochastic) capsules of varying complexity, but a small group possess schizocarpic or indehiscent, fleshy, berry-like fruits. Carpobrotus and Sarcozona are the genera with this later type and together with many of the capsular genera are placed in the subfamily of Ruschioideae on the basis of their parietal placentation (Schwantes, 1957 & 1960). Blake (1969), however, demonstrated that the placentation in these two genera was not only parietal but also partially axile with the placentas extending about half way up the axis. The placing of Carpobrotus and Sarcozona in this sub-family solely upon the parietal placentation is not satisfactory. However, the hybridization of Carpobrotus with Disphyma, described in the present study, does support their retention in the Ruschioideae.

11. Carpobrotus sp. (photo Nick Perrin)


I have no info about the place or time of the year, not sure which of the two species it is (quite hard to distinguish without a flower). It looks like a cultivated plant in some park, possibly in Wellington where Nick lives.

No matter what botanists would consider or how will classify them, these very interesting plants are now part of New Zealand coastal ecosystems and are playing their role in keeping a natural habitat of
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the shores and this will not be taken back for the pride of the New Zealanders. Although Carpobrotus edulis and Carpobrotus aequilaterus are man introduced plants they have found their place here and by creating the new natural hybrids are contributing to the diversity of the New Zealand flora. And if you live in a coastal area just give it a try with cultivating Disphyma australe or even as an annual if you cant offer a frost-free spot, its worth it! References:
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 - http://floraseries.landcareresearch.co.nz ); R. J. Chinnock Studies in Disphyma a Genus Related to Mesembryanthemum, Part 1- A Revision of Disphyma australe (New Zealand Journal of Botany, Vol. 9, 1971); R. J. Chinnock Studies in Disphyma a Genus Related to Mesembryanthemum, Part 2 Infraspecific Division of Disphyma australe and Notes on Australian Species of Disphyma (New Zealand Journal of Botany, Vol. 14, 1976); R. J. Chinnock Natural Hybrids between Disphyma and Carpobrotus (Aizoaceae) in New Zealand (New Zealand Journal of Botany, Vol. 10, 1972); Bruce D. Clarkson A Review of Vegetation Development Following Recent (<450 years) Volcanic Disturbance in North Island, New Zealand (New Zealand Journal of Ecology, Vol. 14, 1990); B. A. Fineran The Vegetation and Flora of Bird Island, Foveaux Strait (New Zealand Journal of Botany, Vol. 4, 1966); A. J. Healy Contributions to a Knowledge of the Adventive Flora of New Zealand Part VIII: The Succulent Element of the Adventive Flora. (Transactions of the Royal Society of New Zealand, Vol. 87, Parts 3 and 4, 1959); P. N. Johnson Vegetation and Flora of Womens Island, Foveaux Strait (New Zealand Journal of Botany, Vol. 14, 1976); New Zealand Plant Conservation Network (2005 - 2007) (www.nzpcn.org.nz ); T. R. Partridge - The Vegetation of Wharekakahu (New Zealand Journal of Botany, Vol. 21, 1983); Nick Perrin Horokaka - The Native Ice Plant (Disphyma australe) (New Zealand Cactus and Succulent Journal, Vol. 49, 1996) Didge Rowe Cultivating Mesembryanthema in New Zealand (New Zealand Cactus and Succulent Journal, Vol. 59 - 64, 2006 / 2007); J. Bastow Wilson & Carol Cullen Coastal Cliff Vegetation of the Catlin Region, Otago, South Island, New Zealand (New Zealand Journal of Botany, Vol. 24, 1986); E. Zimer Disphyma australe ssp. australe (www.cactusi.com Cactus Romania Enciclopedie, 2007).

Further Readings:
Mary E. Gillham - Vegetation of Little Brother Island, Cook Strait (Transactions of the Royal Society of New Zealand, Vol. 88, Part 3, 1960); Mary E. Gillham - Vegetation of New Zealand Shag Colonies (1960) (Transactions of the Royal Society of New Zealand, Vol. 88, Part 3, 1960); P. N. Johnson Vegetation and Flora of the Solander Islands, Southern New Zealand (New Zealand Journal of Botay, Vol. 13, 1975) These articles were not directly referred in this text, nor were any essential or relevant information extracted from, although all three have been consulted. However, these articles are extremely interesting accounts and very technical as well, and useful for understanding the habitat of Disphyma australe or the natural environments down here in New Zealand. All three are available for downloading on the Internet, so that I recommend them to anyone interested by the subject.

Credits & Thanks:


Dag Panco de Grid he knows why! Nick Perrin for the photos and information made available for this article Didge Rowe, Frances Verrity for their help in gathering information Mihai Zimer for helping me out with the English version

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My Notes:
(1) N. E. Brown was the first who used informally the combination Disphyma australe in 1930 and some botanists are accepting his authority. However, although he has set up the characters of the new genus he didnt make any formal combinations. Nick Perrin (1996): The generic name, Disphyma, means two-lumps, and alludes the presence of a two-lobed tubercle at the opening of each seed-containing cell in the fruit. This is the basis on which N. E. Brown separated these plants from other Mesembryanthemums. The specific epithet, australe, means southern. The twin-lumps are lacking in Disphyma australe though, this being the main reason for removing it from the genus. (2) It is not that important if it is or not the same species as the south-african relative; what is important is that it has been recognized as a different species from Disphyma australe that is so for sure! (3) Didge Rowe (in a personal message) is pointing more towards regional differences giving the examples of a pale-pink-flowered intermediate form growing on the west coast beaches near Auckland (Piha, Bethells Beach) and further south (Cooks Strait, near Wellington) one with white flowers and green leaves and another with very red-tinged leaves and darker pink flowers. Chinnock, on the other hand, states that colour variations are quite common within a population. (4) A primary succession is following a natural (such as flooding, landslides or volcanic activity) or man inducted (i.e. deforestation or earthworks of any type) environmental change. Secondary or tertiary successions may occur when the dominant plants are changing themselves the environment enough or are creating the conditions for other plants to take over as dominant form of vegetation. Secondary or tertiary successions have in most of the cases natural causes (we shouldnt blame man for all changes in nature) such as natural ageing or dieback of established vegetation or when those populations are exhausting natural resources or are modifying the substrate in any way which enables new species to colonize. (5) Didge Rowe used to live for many years in Australia and has just recently returned to New Zealand. (6) A study of one of the habitats revealed that Disphyma australe is growing in soils with a pH of 5.0 7.2, the average pH being 6.5; this may not necessary cover the entire range of pH tolerated by the plant, however, the information gives some idea. (7) Reportedly it is quite salt tolerant; seeds are even germinating in water with 1.0% salt concentration, but not with 1.5% salt concentration. (8) Toelken refers now Carpobrotus aequilaterus as an endemic Australian species, different from the South American Carpobrotus chilensis. (9) There are actually different (lets call them) forms of Carpobrotus edulis. The most common form has yellow flowers turning purplish-orange with age, but also known are plants with yellow, pink or pink-orange flower which do not change colour when ageing, which is rather strange as it is pointing to more profound differences rather than a simply flower colour variation. (10) Carpobrotus edulis is actually a very invasive plant, present in (or infesting one would consider) wide areas in California, France and Italy to name just a few having a major impact on the loss of native vegetation forms. For the New Zealand environment it seems to be rather benefic than destructive, although in some cases it may have replaced or limited the spread of native plants. (11) Nick Perrin wrote (in a personal message): Near Wellington airport the D. australe grows close to the sea shore while the Carpobrotus edulis grows further back beside the road. C. edulis flowers that I have seen are very bright yellow and the size of tea cups. In between are numerous hybrid plants with intermediate characteristics between the two species. The hybrids are sterile. Bob Chinnock thought he once saw a hybrid with fruits, but has been unable to find it again. All errors, omissions and misconceptions are mine.

Eduart Zimer, May June 2007

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