Sarcocornia quinqueflora ssp. quinqueflora (Bunge ex Ung.-Sternb.) A. J.

Scott 1977
Abstract A brief account on Sarcocornia quinqueflora ssp. quinqueflora one of the less known succulent halophytes belonging to the family Amaranthaceae with regional Australasian distribution, but having close related species with global distribution. Focus will be set on New Zealand populations and habitats. ***** Sarcocornia quinqueflora ssp. quinqueflora (Bunge ex Ung.-Sternb.) A. J. Scott 1977 is a succulent palustrine plant forming large mounds of articulated stems with leafless appearance. It is still widely referred to the family Chenopodiaceae in most plant classifications (notably Cronquist) but based on recent evidence from molecular phylogenies the APG System (1998) and APG II System (2003) have included this group at subfamily level (Chenopodioideae) in the family of Amaranthaceae [I]. It is therefore not even closely related to any of the major families of succulent plants.

1 - 2. The gap between Motutapu and Rangitoto can be crossed by foot at low tide, but the shallow and usually very calm waters of the northern part of the Islington Bay (called also Easy Bay as it used to be a resting place for drunken sailors) and Gardiner Gap are ideal for salt marsh vegetation (left).The gap between Motutapu and Rangitoto, while the tide is picking up (right).

The succulent plants belonging to Chenopodioideae (Chenopodiaceae) have various vernacular names such as glasswort or samphires. This particular plant I am writing about was formerly known as Salicornia australis Solander ex G. Bentham 1870 (1) and Arthrocnemum australasicum (Moquin Tandon) Moss and is a New Zealand native glasswort growing sometimes quite abundant on salt marshes, sandy beaches, and rock platforms with some interruptions on both coasts of the North Island and mainly on the eastern coasts of the South Island, and on the northern parts of the Stewart Island as well. The plant has actually a quite wide regional distribution including Australia, Chatham Islands, Norfolk Islands and New Caledonia. A distinct subspecies Sarcocornia quinqueflora ssp. tasmanica inhabits the Australian state of Victoria and the Tasman Island shores and another closely related species Sarcocornia blackiana is widespread in Southern and Western Australia; the later one is characterized by a thicker fruiting spike which is the main noticeable difference. Page 1

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3 - 4. Sarcocornia quinqueflora with Agropyron junceiforme on lava blocks in Rangitoto. This particular plant grows between lava blocks taking advantage of the very tight space where shingle can accumulate. The usually substrate is loose and consists of very good draining shingle or shingle mixed with sand and crushed shells (left). Sarcocornia quinqueflora stems of a plant from Rangitoto (right).

5 - 7. Flowering plants in Rangitoto. The flowers are really tiny and nondescriptive, and probably of no interest whatsoever for any amateur. I havent insisted in close-ups as well (above left). Sarcocornia quinqueflora growing on shingle between lava blocks in Rangitoto. The substrate (and the debris collected by the packed stems) is better visible in this picture (above right). Flowering plants in Rangitoto (left).

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It grows abundantly from the north-eastern Australian coasts all the way down to the south-east and also in the south-western regions; it is also widespread in New Zealand, more scattered and in smaller groups in North Island but forming on occasion massive populations in South Island where it is still very widespread despite of the constant decline of the salt marsh areas its main habitat type. It is usually restricted to the shore line, but it can sometimes grow in inland localities (two inland populations are known in New Zealand in Central Otago, and much more in Australia); however, it grows often along the salt marshes, at the edges of inundated areas, mudflats, lagoons, estuaries and the tidal river banks through the inland sometimes reaching 10 15 km from the sea shore. Pure inland populations are rather rare and restricted to high salinity soils or moving sand dunes. However, the plant occupies also a secondary habitat type, quite different from the tidal areas, consisting of coralline, sandstone or porous volcanic rocks, platforms and cliffs, very close to the waterline, in most of the cases within the splashzone, but sometimes growing even 10 15 m above sea level. The main difference consists not necessarily in the absence of salt, as there still is a strong marine exposure with all that comes with it including strong winds and misted salty seawater, but the absence of partly submerged or waterlogged conditions; instead the plant has to gap periods of bone-dry substrate. This secondary habitat type provides smaller plants, with a different growth form. However, the plants tend to occur in the proximity of the sea not only because they have adapted to high salinity levels, but because they can not face the competition of upslope plants with moderate resistance to salt such as Disphyma australe or Crassula moschata to name the succulent species, or such as Hebe elliptica and Poa astonii capable to grow through most of the vegetation sequence and having therefore a wider distribution (Wilson & Cullen, 1986).

8 - 9. Sarcocornia quinqueflora with Agropyron junceiforme on shingle between lava blocks in Rangitoto. The mangroves (Avicennia resinifera) seen in the background will take possibly over in time. The places where usually Sarcocornia quinqueflora grows here are partly submerged during high tide (left).A site with a typical salt marsh aspect in Rangitoto. Again you can see in the background young mangroves ready to take over the habitat.

The plant is virtually unknown from cultivation and also did not have any economical or culinary importance as its European relative Salicornia europea the original glasswort - used to have (2); it is not known for Sarcocornia quinqueflora or any other related plant to have been used as food or for other particular purpose by the New Zealand and Australian native populations. However, ecologists highly value this plant is for providing groundcover and preventing soil erosion in salt scalded areas. It has little to no economical importance though because of the distribution areas; even in inland localities it does not provide a grazing resource as it usually does not recover readily from defoliation, trampling or fire. Page 3

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10 - 11. Sarcocornia quinqueflora in Rangitoto (left). Russell Bay of Islands (Northland) habitat just few hundred meters north of the wharf. You can see the tide line at the bottom of the rocks. As idyllic as it may look, this habitat with a western aspect has strong marine exposure. In the lower parts of the rocks you can see an almost continuous strip of Crassula multicava.

Contrary to the most of the glassworts, having an annual habit or turning under circumstances to biannual or short lived perennial habit, Sarcocornia quinqueflora ssp. quinqueflora is a typical perennial plant, with distinct growth and resting periods. It has fleshy perennial stems spreading first horizontally then turning upwards, forming in some cases a bushy plant, although in most cases it forms just a 15 30 cm high dense mat. In sheltered positions plants may grow up to 60 cm high, usually single plants growing in not too exposed places are growing bigger and taller. Sometimes in mostly procumbent plants the stems may reach even 90 cm in length and try to fix themselves by shooting adventive roots at the nodes. Branching is usually opposite. The stems are mostly cylindrical, sometimes to narrowly obovoid, green to gray-green when young, with a waxy appearance, later turning from yellow to orange to red and brown, very succulent, later becoming corky usually in the second year. F. W. Cooke (1912), the author of a very early but highly interesting and detailed paper on this plant, wrote: As winter advances, as a rule, all the branches formed in spring assume a different appearance. The succulent tissue becomes withered and turns brown, the free portion of the leaves of each internode surrounding the base of the internode above like a collar. This brown portion finally falls off, or, if the plant is at the waters edge, is soon washed off, and the branches appear green again. They are, however, much smaller, having lost all palisade and aqueous tissue. () This withering of tissue is due to the formation of cork. The inner layer of pericycle, which is now several cells thick, gives rise to phellogen. Cork tissue and phelloderm are formed in the usual way. The stems have usually 2 4 mm diameter, but quite frequently up to 5 6 mm (and on occasion even more in older plants) and the articulated internodes are 5 15 mm long. The stems have the normal apical growth but may also increase greatly in thickness over time due to secondary growth. In case of older plants the diameter of the stems may reach 10 12 mm. The stems are jointed, and are bearing the characteristic much reduced leaves and appear to be leafless at first sight the small succulent lobes at the apex of the internodes are the leaves, mostly minute but sometimes getting larger. The dorsal surface of the leaves is concave, the ventral being convex; the leaves are populated with numerous stomata on the ventral surface and not at all on the dorsal surface. The leaf has a very uncommon anatomy F. W. Cooke (1912): The leaf margins are colourless, since, being only two cells in width, there is no palisade tissue developed between the dorsal and the ventral epidermis. Stomata are formed usually starting with the 4th leaf from the apex and are situated above the palisade tissue which is quite unusual for most of the plants but common for plants of this type. Plants have sometimes a basal concentration of leaves, but this is not a rule. Photosynthesis takes place in leaves and stems as well. The Page 4

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colour of the stems is varying sometimes in a very strange way - F. W. Cooke (1912): A curious feature is shown in some stems, especially in those on which an inflorescence has been borne. The internode nearest the stem is withered and brown, several adjoining it are green and succulent, then again there are several brown and withered internodes, then either the succulent apex or the withered remains of the inflorescence. Colouring always starts in the first internode from the base of the plant and is depending on the location, the more exposed plants or plants growing on rocks are inclined to turn red earlier; this colour doesnt stay too long as the stems are getting brown soon. The apparent withering of the internodes is not a sign that the plant will die, plants that are dieing prematurely do not usually turn red. The withered appearance is stronger in autumn when many plants from exposed positions appear to die down, just to sprout out again in late winter; in other cases the plants remain more or less green through the year.

12 - 13. Russell Bay of Islands (Northland) habitat just few hundred meters north of the wharf. A slightly more exposed rock with a small Sarcocornia quinqueflora tussock in the centre (left). One the plants in Russell Bay of Islands close to the wharf, growing on the rock pictured in photo No. 12 (below).

Sarcocornia quinqueflora ssp. quinqueflora flowers in December to March (during the austral summer) and very probably the flowers are wind pollinated at least no insects were observed working on those flowers and no nectar glands are present. Most of the branches (not necessarily all) are producing insignificant flowers in the apical region, in the axils of the leaves, and can form sometimes an almost complete ring. The number of the flowers in each axil is usually 5 hence the name quinqueflora but it may vary from 5 10 (F. W. Cooke, 1912). The circular seeds are 1.5 mm small and covered with acute hairs. Seed dispersal is definitely by wind; however, I think that the hairy outlay allows for enough floatability to be dispersed (at least for short distances) by sea currents and waves. I am making just an assumption in this case, but based on similar dispersal means of another succulent halophyte naturalized here in New Zealand Cakile maritima.

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The plant is self-compatible but H. E. Connor (1984) has no doubt that intra-plant pollination occurs in nature. However, the fertility is very high, the set-seed being in average 94.5% for fresh seed. There are three types of plants plants bearing hermaphrodite flowers, plants bearing functionally male flowers and plants bearing functionally female flowers. It is very interesting that these types of plants do not occur usually at the same sites. Plants bearing male flowers are not known in New Zealand and the very most of the populations consist of hermaphrodite plants only. In just three areas of New Zealand mixed populations (hermaphrodite and female plants) are known. Female plants are usually less frequent; as H. E. Connor (1984) shows in several populations from Golden Bay the female plants have an average frequency of 32% (varying from 12 53% within different sub-populations); in Tasman Bay female plants are 36% from the total (varying from 19 41% within different sub-populations); in Otago Peninsula populations on the other hand the frequency of the female plants is way less no more than 5 10%, possibly even lower at Hooper Inlet. Despite of this, except for the sand habitat, no other ecology variations could be established between the hermaphrodite and female plants. In a fourth small population considered a mixed one, east of Picton (north of the South Island) the presence of female plants couldnt be confirmed by H. E. Connor; reportedly 25 hermaphrodite plants and 2 female plants were observed at an earlier date.

14 - 15. Sarcocornia quinqueflora at Tahuna Torea (Auckland). The other succulent plant seen growing here was Carpobrotus edulis, the yellow flowering form (left). Sarcocornia quinqueflora at Tahuna Torea (right).

Sarcocornia quinqueflora ssp. quinqueflora has a quite high resistance to salt; adult plants grow fine up to a concentration of 3.0 3.5% salt. On the other hand the germination is poor or impossible if the salinity is not significant lower (it is quite high under 1%, declining fast between 1.0 1.5% and is only sporadic at much higher concentrations). The salt marsh vegetation consists of annual and perennial plants (some of them succulent) and each group has developed a different survival strategy. T. R. Partridge & J. B. Wilson (1987) reveal in their study a very interesting aspect the seeds of the annual plants (and of the short-lived perennials as well) are not very sensitive to high salinity and are germinating readily in these conditions; the seeds of the perennials on the contrary are quite salt sensitive and can germinate only in salinities much lower than those that can be tolerated by adult plants. The higher the salt tolerance is in adult plants, the higher is the sensitivity of the seeds. These differences between the annuals and perennials are highly influencing their habitat and colonizing ability. The annuals are producing usually seed in big quantities and these are germinating readily even in quite hostile conditions this feature allows them to act as aggressive colonizers on the lagoon edges, in disturbed marsh or anywhere the sand and mud becomes for some reason exposed. This is explained by Page 6

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the fact that their seed germinates independently on the salinity level, mature plants can tolerate high salinity when this occurs, and, not of least importance, due to their programmed vegetation pattern they have to complete rapidly their life cycle in a given time and in a hostile or unpredictable environment. A different path has been chosen by the perennials, as ideal conditions for germination are exceptionally rare in salt marshes and similar habitats. The perennials have so to speak all the time to wait until ideal conditions are met (especially low salinity due to heavy rains, floods, rising or draining of the mud or sand flats, etc.) in order to allow germination and establishing of the seedlings. Annual plants cannot afford to do so.

16. Tahuna Torea (Auckland) at low tide.

As earlier mentioned there are two types of habitats quite different from each other - in which Sarcocornia quinqueflora ssp. quinqueflora may be seen the mostly wet salt marsh habitat type and the dry sand dunes or rocks and cliffs habitat type; each of them producing plants with different growth types and characteristics. But first of all and a bit surprisingly there is actually no stringent need of a salt laden habitat for these plants to thrive. J. Bastow Wilson & Carol Cullen (1986) consider that extreme halophytes (and they mention Sarcocornia quinqueflora) have a requirement of salt just in order to have a maximal growth rate. In their study they show that these plants tend to occur in lower parts of the cliffs, with higher salinity although the salt requirements are not absolute; they can grow at two thirds of their maximal rate in fresh water . It is just their adaptive and specialized behaviour that makes possible to survive such adverse conditions and probably the inability to face competition in more favourable environments and this keeps them away from having a wide distribution range outside their usual habitats. In fact the plant does not take up significant amounts of water and it is not longing at all for the high-tide. The roots are more or less sealed and do not perform their usual functions in full unless the water has a lower salinity (3) as it can happen every now and then during heavy rains (associated with low-tide) or

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floods. Even waterlogged it keeps its xerophytic character as its survival does not depend on a permanent source of moisture.
17 - 19. Salt marsh habitat dominated by Avicennia resinifera at Pahurehure (Auckland); there is no place here anymore for Sarcocornia quinqueflora, the wooden mangrove being the stronger succession plant (left). You can see how moist it can get at times on the vertical cliffs due to the draining fresh water pouring from above and the sea at its feet in Raglan (Waikato) (below left). Sarcocornia quinqueflora - a plant from Raglan (Waikato) struggling with extreme moist conditions this time it is fresh water (below right).

The main habitat type of this plant is the salt marsh where Sarcocornia quinqueflora ssp. quinqueflora usually grows on loose shingle and is less common on fine and sandy or compacted sedimentary substrates; it sounds simple, but it is in fact a very complex and highly variable environment. For a relatively small country New Zealand has a very varied shore line and there are also some other factors making the regional differences even deeper. First of all there is a big difference between the eastern coastline (dryer) and the western coastline (wet) due to a specific dominant weather system. This is of great importance as we are dealing generally with succulent flora. In the North Island there is warm enough to make the establishing of significant populations possible, but in the South Island due to a significant cooler climate associated locally with extreme high rainfalls it is virtually impossible for the succulent flora (including Sarcocornia quinqueflora ssp. quinqueflora) to establish significant wild populations on the western coast. On the other hand the regions in which estuaries are numerous and salt marshes are present reach from Northland to Southland but the conditions and the eco-types are rather different. In the northern parts the populations of Sarcocornia quinqueflora ssp. quinqueflora are smaller Page 8

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and more restricted due to the massive presence of Avicennia resinifera , the wooden mangrove, a plant with rather subtropical needs (restricted to the North Island) and another tall plant Juncus maritimus var. australiensis, which has its southern limits of distribution just north of coastal Otago. These two plants are very important competitors in the waterlogged areas and are both directly influencing the natural habitat. This makes possible that from Otago southwards the vegetation patterns of the estuaries and salt marshes are different, with predominant low vegetation (usually less than 10 cm) and with very restricted taller vascular plants. Due to this the southern half of the South Island populations are generally more consistent and larger; in some places even becoming the dominant form of vegetation, usually very abundant in the lower marshes in waterlogged conditions (some of the companion plants are Puccinellia novae-zelandiae, Samolus repens, Atriplex prostata, Triglochin striatum) in most of the cases having just little fresh water available, mixed with the tidal sea water. On the contrary North Island populations seem to be more fragmented, consisting more of relative few plants scattered in exposed tidal areas, on shingle, near beaches or cliffs and rocky platforms with shallow soils accumulated in pockets, on cyclic drained mudflats where mangroves have not established significant populations.

20 - 21. Details of plants growing in Yankee Wharf (Rangitoto).

Salt marshes are extremely complex eco-systems with each location having actually its own character. T. R. Partridge & J. B. Wilson (1988) make an excellent account of a southern-type habitat, where wooden mangroves are absent: A notable feature of the individual marsh classifications () was the abundant, species-poor, and repetitive lower marsh. () Within each marsh the composition of the lower marsh is quite consistent, but the species mix varies from one marsh to another. () There is no consistent grouping to be found. At Karitane North, the species associated with Sarcocornia quinqueflora is a different grass, Puccinellia fasciculata, perhaps because the marsh is sandy and prone to drying when exposed. () Another species, the taller Puccinellia novae-zealandiae, seems restricted to sheltered Page 9

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muddy marshes at Cherry Farm and Papatowai and occurs as a distinct but narrow band below the Sarcocornia quinqueflora. Included in lower marshes are raised sandy margins that often occur around the seaward edge of exposed maritime marshes () Wind-generated waves pile up sand and splash sea water on those margins. Sarcocornia quinqueflora is common, but the species indicative of such wavesplash conditions is another succulent chenopod, Suadea novae-zealandae. Such areas become extremely saline when water evaporates, concentrating the salt. In fact more or less erratic combinations of very different factors (waves, evaporation, heavy rains, fresh water floods, winds and their direction, tides, etc.) may drop the salinity level to less than 1% or boost it to 5% in (and for) a very short time. Erratic and temporary changes are part of the eco-system, but long time and consistent changes especially of the salinity levels may alter massively the habitat. Due to this fact salt marshes are in a very fragile balance and extremely exposed to mans interference (4). There is also another difference given by the lagoonal / estuarine character of the salt marsh. Although many factors are involved, the main difference is that estuarine salt marshes are object of a regular tidal flooding (and subsequently it never gets too saline, it never gets too fresh, it never dries out completely), but the lagoonal type has irregular and often sustained flooding of fresh, brackish or sea water, regulated by the frequency and grade of the opening of the sand bar at the river mouth. T. R. Partridge & J. B. Wilson (1988) give an extreme example with Kaikorai (Otago) where in some places due to this factor the plant communities bear little resemblance with the typical salt marsh vegetation, where some species of the lower marshes have been replaced by short-lived perennial opportunists such as Cotula coronopifolia and Spergularia media, invading exposed banks, or Isolepis cernua, which can even survive longer periods of subsequent inundation. Annuals and short-lived perennials are perfectly suited for this kind of irregular flooding schedule. The most impressive salt marsh population I have seen here in New Zealand was that of Rangitoto Island (Hauraki Gulf) on the low and rocky coast, between lava blocks and shingle, in places where the seabed was exposed during low tide and barely submerged on high tide. The nice clumps of the salt marsh at Tahuna Torea (Auckland) where also very interesting, but unfortunately I havent visited yet the extensive populations of the South Island, nor the Napier salt meadow where the most extensive population of the North Island resides.

22. The largest population I have seen so far in a shallow bay close to the Rangitoto wharf in the southern part of the island. During the high tide most of the plants are waterlogged.

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The dry habitat of the sand dunes or rocky slopes and coastal cliffs produces different plants, with smaller and with less upright growing stems, but with a rather contorted and irregular growth. The plants growing on dry substrate have not the mostly uniform gray-green colour of those growing in salt marshes, bearing little resemblance for an inexperienced observer. It was with big surprise to see few patches of miniature plants growing on almost vertical cliffs at 8 10 meters above the sea level at Muriwai, few kilometers out of Auckland at the Tasman Sea, forming a low mat probably not higher than 10 - 15 cm, or at Russell (Bay of Islands), growing on a low sandstone steep, just above the tidal line. But probably the most remarkable plant seen was that of Devonport wharf (Auckland) a single non-descriptive small plant growing in the cracks of a concrete block in the splash zone; this tiny poor plant, completely isolated and fighting for its survival made me think that seed dispersal by combined action of sea currents, tides and waves must be also considered, as no other obvious seed source was available nearby. What could have been the reason for this plant to choose the evolutionary pathway of true succulence when it has a lot of water available at least in its typical salt marsh habitat - at any time? As any other glasswort such as Salicornia virginica or Salicornia europea, this plant is designed to minimize water loss. The leaves have been reduced to a minimum (in most plants on the brink of disappearance) to avoid transpiration, same with the flowers; stems have also a photosynthesis function. The epidermis of the plant has a waxy appearance and is keeping the moisture inside the plants; un-rooted plants may resist several weeks, even months if not exposed to extreme conditions, without any sign of significant dehydration. The stems have also the ability to store water like any other succulent plant; even very old stems do not become completely corky or wooden, and there is still succulent tissue present. F. W. Cooke (1912) described the aqueous tissues, formed of large cells with thin walls apparently specialized in water storage as these cells are filled with water at any time of the year (at least in plants not dieing back) and large intercellular spaces suggesting that this makes possible for the plant to take up big quantities of water in a short time if needed and also to store it. Sarcocornia quinqueflora ssp. quinqueflora loves the waterlogged conditions and is the only habitat type in which it may sometimes become the dominant form of vegetation, but it is not the only place this plant can be found. I would say that it is probably its original habitat, but this plant is a survivor and probably has the potential to move to other habitat types if competition easies. I would say, half serious, that it is an open minded being exploring other opportunities as well. Lets go straight to the opposite situation the steep rocks and cliffs. This is a very dry and hostile environment (not to speak of the lack of any organic matter) and even if rain is abundant all the water flows away, the little moisture retained by the cracks or within the porous texture of some rocks does not last very long. Under normal circumstances in New Zealand conditions with no other rains following, but strong winds and sunshine, it would dry out completely in a matter of days. The plant has just a limited time in which it can absorb the available moisture. Sarcocornia quinqueflora ssp. quinqueflora and other similar plants are xerophytic-halophyte plants, being able to survive in reasonable dry environments with high salinity. It does not occur in big numbers in this kind of habitat (as it does in the wet) but it is a frequent presence.

Referred Literature:
ABRS Flora of Australia Online (Data derived from Flora of Australia Volume 49 (1994), a product of ABRS, Commonwealth of Australia); 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 ); 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);

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H. E. Connor - Gynodioecism in Sarcocornia quinqueflora (New Zealand Journal of Botany, Vol. 22, 1984); F. W. Cooke Observations on Salicornia australis (Transactions of the Royal Society of New Zealand, Vol. 44, 1912); J. Gathe Sarcocornia (contribution for the electronic version of The Western Australia Flora (2000 - 2007), http://florabase.calm.wa.gov.au ); T. R. Partridge & J. B. Wilson Germination in relation to Salinity in Some Plants of Salt Marshes in Otago, New Zealand (New Zealand Journal of Botany, Vol. 25, 1987); T. R. Partridge & J. B. Wilson Vegetation Patterns in Salt Marshes of Otago, New Zealand (New Zealand Journal of Botany, Vol. 26, 1988); G. Sainty Sarcocornia quinqueflora (2006); E. Zimer - Succulent plants from down under Sarcocornia quinqueflora an odd succulent halophyte (2007, http://eduart.page.tl/).

Further Readings:
R.W.G. Carter & Bill Carter - Coastal Environments An Introduction to the Physical, Ecological and Cultural Systems of Coastlines (1988); John R. Packham & Arthur John Willis Ecology of Dunes, Salt Marsh and Shingle (1997); T. R. Partridge - The Vegetation of Wharekakahu (New Zealand Journal of Botany, Vol. 21, 1983).

My Notes:
(1) The New Zealand plants are mostly referred as Salicornia australis Solander ex Forst f. 1786, but this is a nomen nudum. H. H. Allen considers that the New Zealand plants might be not conspecific with the Australian plants but also admits that this matter has not been studied adequately. He is referring to McCanns studies back in 1952 who was revealing floral variances from the Australian populations (the presence of two minute orbicular deeply concaved, membranous floral lobes, () perhaps corresponding to a corolla ) which may support the segregation as a distinct subspecies or form. However, new studies of fresh material have not confirmed so far the variances so that the possibility of McCann dealing with abnormal material cannot be excluded. The vernacular name glasswort was applied originally to the widespread Salicornia europea (and later to other Salicornia species) as this plant was used in the early times of glass production in Western Europe, due to the high soda content of the ashes. It was also widely used in medieval and even post-medieval times as a survival food so to speak, later named the poor mans asparagus. Nowadays it has become an exquisite main ingredient for gourmet restaurants especially on the eastern coasts of England. Reportedly it has a quite distinct salty and slightly astringent taste. The root system is not pumping continuously salty water into the plants body because it would end up having a deadly concentration of salt. On the other hand a certain concentration of salt is necessary for the plant in order to survive. Usually the plants body is wetted frequently with salty sea water (or is even partly submerged for some time) and if the inner sap is not salty enough due to the phenomenon of osmosis water would be extracted from the plants body in order to balance the external concentration. Thats why the plant has an inner salinity high enough to avoid this from happening, but still it has to limit carefully the amount of water pumped into its body by the root system, unless it is not fresh or brackish water. Cultural modifications have had a great impact as it encourages the encroaching of land development on to the marsh, heavy grazing in the upper zones is also creating all the conditions for invasive plants to settle and replace the typical (and sometimes endemic or endangered) vegetation. On occasion these changes may affect even the sand-binder vegetation of the sand dunes and as a consequence it may induce even more instability. Stable sand dunes (former beach fronts) have been also modified to farmland especially in the years of the European settlement boom of the end of the 19th century and oversown with pasture grasses or other agricultural plants replacing partly or totally the native vegetation.

(2)

(3)

(4)

Note of the German translator (Martin Staege):

[I]. This very interesting subject is still under debate. See e.g. K.A. Shepherd (2008): Consensus or complacency? A discussion of the proposed new collection sequence at the WA Herbarium. Australian Systematic Botany Society Newsletter 135:5-8.

Eduart Zimer, November 2008 eduartzimer@yahoo.co.nz http://eduart.page.tl/Home.htm Page 12

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