A conservation overview of Australian non-marine lichens, bryophytes, algae and fungi
G.A.M. Scott, T.J. Entwisle, T.W. May & G.N. Stevens
Environment Australia, May 1997
ISBN 0 6422 1399 2
Bryophytes
George A. M. Scott
3.1. Significance and reasons for conservation
Conservation of bryophytes as such – that is, in contrast to the conservation of plants in general, which may include bryophytes – is a matter which is just beginning to concern governments and, at that, in only a handful of countries. A recent symposium (Hedenäs & Soderström 1992) which forms a useful starting point and stimulus, dealt more with needs and problems than with active achievement, but a World Red List of endangered bryophytes (Geissler et al. 1993) is being compiled and there are lists for individual regions (e.g. Gradstein 1992). The alarming actual and potential consequences of acid rain have stimulated more activity in Europe than elsewhere and it seems probable that the decay of Sphagnum bogs there will make Sphagnum the front-runner in bryophyte conservation. Already the UK has investigated guidelines for bryophyte conservation (Hodgetts 1992) but there are also strong movements in Scandinavian countries and elsewhere in Europe, in north America, and the beginnings of concern in Japan (Takakai & Ando 1975). A realisation of the significance of bryophytes in ecosystems and appreciation of the needs for conservation are slowly gathering force, but it would not be realistic to expect any progress except in countries where there has first been considerable progress in the conservation of vascular plants and where there are also a significant number of bryologists and considerable bryological activity and knowledge. In Australia and New Zealand these criteria are met – more or less – and the time is ripe for bryophyte conservation to be investigated seriously.
The significance of bryophytes in Australia is of two kinds: natural (ecological and aesthetic) and technological. Their roles in ecological processes have been discussed in a number of papers (Scott 1982b; Moore & Scott 1979). They form an important part of the biodiversity of Australia, and in some communities such as wet forests and mallee can sometimes contribute more to biodiversity than any other group of plants.
The aesthetic significance of bryophytes is easy to state but hard to prove. Without them, rainforest would be merely wet, not lush; bogs, if they could exist at all, would be merely quagmires; the mallee would be sandy desert; and mountain rocks would be barren. Much of the attractiveness of these habitats would be gone. Mosses and liverworts, as well as being a key part of the mechanism that keeps many habitats healthy, are also an important part of the ambience that keeps them attractive.
3.1.1. Ecological
The significance of bryophytes in Australian ecosystems depends on four properties: their ability to stabilise soils, to trap and hold moisture, to exchange cations, and to tolerate desiccation. These attributes are enhanced by their ability to spread by both branching growth with subsequent decay of connections, and by fragmentation with subsequent development of even minute fragments into full individuals (i.e. high totipotency). The habitats in which their roles are of greatest significance are: seashore and inland sand dunes, semi-arid soil crusts, wet forests, heath/bog lands, and clay banks; possibly also rock outcrops in both lowland and highland regions, but the significance there is less clear.
Sand dunes
Bryophytes are rarely, if ever, initial colonisers of mobile sand, but once the initial sand movement is locally stopped (usually by algae or angiosperms), bryophytes – mainly mosses – take over as a major stabilising force, especially a few dominant, fast-growing species such as Barbula (Tortella) calycina, B. torquata, Tortula princeps (recently distinguished as T. antarctica), Triquetrella papillata (Moore & Scott 1979). Once established, they are capable of withstanding periods of burial by fresh sand (provided it is not too deep-not more than 4 cm), with outward flexing of leaves in the change from dry to wet positions, allowing them to 'swim' up through the overlying sand, packing it down around them where subsequent rhizoid production can bind it together.
Soil crusts
The soils of much of semi-arid Australia are held in place by crusts predominantly composed of lichens, algae and bryophytes (and sometimes also fungal filaments), preventing the soil from blowing away. It was the removal of this crust by ploughing which brought about the classic topsoil dust-clouds of the Mallee. It is only in recent years that the importance of these soil crusts has begun to be recognised, both in Australia and overseas, and conservative cultivation practised (Scott 1982b). The exact composition of the crusts, the proportions of lichens, bryophytes and algae, reflect local conditions of moisture, nutrients, litter, stability, and time since colonisation, as well as chance historical factors and no doubt the frequency of fire, but insufficient work has been done for us to interpret the significance of the different crust compositions. Undoubtedly many of them are very long-lived – certainly decades and probably centuries – and potentially contain the history of their development, but we are not yet in a position to read it. This role in holding soils in place is of very great importance, although scarcely appreciated.
Wet forest
Particularly in cool- and warm-temperate rainforests, much of the trunks, branches and even twigs of trees and shrubs is coated with bryophytes and a substantial proportion of the ground may also be covered. These bryophytes act in at least three ways: by slowing down and delaying run-off during rain, and hence diminishing erosion; by retaining moisture after rain, and hence maintaining the high humidity regime within rainforests; and (although this is only inferred and not directly observed) by filtering both through-fall and run-off through the bryophyte carpet's cation-exchange system. The significance of this in forest ecosystems has not been established in Australia although it has been overseas (Bates 1990).
A more specialised niche is occupied by epiphylls, which are bryophytes (and lichens) growing on the leaves of other plants in rainforest. Scientific research on the nature of this relationship is just beginning (Olarinmoye 1974), but already it is clear that nutrient flux is involved as well as surface phenomena (Coley et al. 1993). The ecological significance is not clear, but the scientific interest is very high and epiphylls are also one of the most reliable of all indicators of humidity regimes at the higher end of the scale.
Sphagnum bogs and heaths
Sphagnum has two physiological properties of great ecological importance: it has been shown to act as a cation-exchange substance by virtue of the chemical composition of its cell walls (Clymo 1963, 1982) and it has, because of its unique structure, a very great water-holding capacity, holding up to 20–25 times its own weight of water – hence its use as an absorbent and in horticulture. The significance of Sphagnum in the hydrology of high-country boglands is well established. Reports of erosion of subalpine Sphagnum bogs as a result of intensive cattle grazing have been grounds for public concern in Victoria and New South Wales, and would apply equally in Tasmania. The effect of cation-exchange in nutrient cycling in upland bogs has not been investigated but is likely to be significant.
Clay banks
When clay-rich soil has been laid bare either by natural processes such as landslides or by road-making, colonisation and stabilisation are substantially by the agency of bryophytes, particularly in wetter, forested regions. The soil surface rapidly becomes bound together by both rhizoid production and by co-planar branching of prostrate stems, and is thus shielded from further erosion. This process fails only where there is inadequate moisture (and in such cases erosion tends to be minimal anyway) or where exposed to full sun, in which case colonisation tends to be much slower. The process of colonisation has not been studied in detail but it is a normal feature of forests from Tasmania to far north Queensland, and in south-west Western Australia.
Rock outcrops
Bryophytes – both mosses and liverworts – are characteristic colonisers of bare rock in both lowland and upland regions, usually occupying different (probably periodically moister) micro-habitats from lichens which also colonise similar rocks. Apart from their familiar function in delaying and slowing run-off, mosses are often held to initiate colonisation of bare rock surfaces, leading on, through a classic lithosere, to final complete colonisation by flowering plants. Bryophytes certainly are major colonisers in the build-up of humus on rocks such as the top of granite outcrops in Western Australia, but there is little evidence that this leads on to later stages of succession except where the rocks are so low that colonisation can in any case encroach from the sides. In other cases, in fact, such as the sides of outcrops, the succession is clearly cyclical with a build-up of bryophytes and/or lichen colonies holding too much moisture for the strength of the anchoring rhizoids (and rhizines) and the whole community sloughs off, leaving nearly bare rock to start again. This process undoubtedly leads to humus production but any effect it might have in breaking down rocks must be measured over a very long timescale, although the role of lichens (as distinct from bryophytes) in rock erosion is well documented.
3.1.2. Practical benefits
Commercial
Commercially, bryophytes are one of the least exploited groups of plants. Only Sphagnum has widespread economic significance, although there are others of less importance (Ando & Matsuo 1984); the cultivation of bryophytes in gardens has a long history but apparently is extensively practised only in Japan. Their phytochemical potential for medicinal and practical use has scarcely been looked at, let alone explored. Because of their antiquity as a group (going back at least to the Devonian) their genetic resources are likely to be very different from other groups and hence to have unique potential, as yet untapped. i.e. there is a strong case for phylogenetic conservation (see Section 1.2). The walls of liverwort cells, for example, differ greatly from those of higher plants in having a much greater resistance to chemical reagents and decay as well as animal attack – a feature of obvious implications for genetic engineering.
Scientific
Bryophytes have a unique combination of comparably sized but structurally very different haploid and diploid generations, easily cultivated and ideal for investigating the effects of developmental environment versus genetic control in determining morphology. Their potential for investigating the gap in knowledge between genotype and phenotype, between the genetic blueprint and the resulting organism, i.e. microdevelopmental control, is unsurpassed, although only slowly achieving recognition. Because of their anatomical simplicity they are likewise an important tool in the investigation of basic physiological processes such as photosynthesis.
As well as their inherent interest for their remarkable range of form, beauty and elegant adaptive and morphological features, they occupy a pivotal position in the evolutionary switch from gametophyte-dominance to sporophyte-dominance, and in the concomitant switch from primordial life in water to subsequent life on land.
3.1.3. Conservation
The reasons why it is important to conserve bryophytes as a group are straightforward: their ecological importance is critical in many habitats and their scientific potential is very great and still untapped. The reasons for conserving individual rare or endangered species are more tenuous but are the same as with species of any other plant or animal group: the world would be a poorer place with the loss of any of its components, and we are responsible for all the forms of life of which we have stewardship. In the last analysis it is a moral imperative.
3.2. History of conservation
At present, with the exception of all species of Sphagnum in New South Wales and the moss Pleurophascum occidentale in Western Australia, no individual species of bryophyte is protected in any State in Australia, although where particular communities are given protection, the bryophytes within them will be incidentally protected also. In general, protection given to 'flora' will include bryophytes (and other cryptogams). There has been no pressure in any State for the conservation of bryophytes per se although there have been recommendations to reserve Sphagnum-rich areas in Tasmania (Whinam et al. 1989). The present conservation position in the different legislatures is summarised in the Introduction, Section 1.2.
3.3. Present conservation status
3.3.1 Taxonomic scope
The bryophytes comprise two or more groups (depending on taxonomic opinion); for simplicity they may be considered as two main groups – mosses, and liverworts or hepatics (including hornworts). In Australia the numbers of species are not known with any certainty. Current estimates of 400–500 extant liverworts (Scott 1982a) and more than 600–700 mosses (Scott 1982a) are probably conservative (see below). Taxonomic knowledge of the Australian flora has been appraised by Scott (1982a) and is at a stage where the primary survey work, although far from comprehensive, is adequate for the production of a provisional flora and checklist for most States, and work is proceeding on individual genera.
There are already checklists and quasi-checklists (although some are already outdated) for:
- Victoria
Mosses & liverworts. Cropper et al. (1991). - Queensland
Liverworts (far from complete). Windolf (1987).
Mosses. Stone (in preparation). - New South Wales
Mosses. Ramsay (1984a). - Lord Howe Island
Mosses. Ramsay (1984b). - Australian Capital Territory
Mosses. Ramsay & Streimann (1984). - Tasmania
Mosses. Dalton et al. (1991).
Liverworts. Ratkowsky (1987).
Handbook of liverworts in preparation (Engel). - Northern Territory
Mosses. Catcheside & Stone (1988). - Western Australia
Mosses. Stoneburner et al. (1993). - South Australia
Mosses. No checklist.
Handbook of Mosses. Catcheside (1980)
Liverworts. No checklist.
More general handbooks cover the southern regions: Scott & Stone (1976) for mosses and Scott (1985) for liverworts. Catcheside (1980), although not exhaustive, is effective for the dry regions of all States and Territories. There are checklists for the whole of Australia of mosses (Streimann & Curnow 1989) and liverworts (Scott & Bradshaw 1986).
It seems probable that the planned volumes on mosses and liverworts for the Flora of Australia will be near publication or at least in progress by the end of this century. Because of the stage that taxonomic work has reached, the expertise on individual genera is now mainly with overseas experts. In Australia there is a small group of workers who keep in touch regularly through the Australian Bryological Newsletter and an annual field meeting with the counterpart New Zealand group. Most active workers in Australia are retired and their potential successors have been inhibited by an almost total lack of employment prospects. Only two Australian herbaria current employ bryologists (Australian National Herbarium: Mr H. Streimann; State Herbarium of South Australia: Dr B.G. Bell), although there are some competent amateur and retired assistants. In total only half a dozen are employed as bryologists.
Estimates of the numbers of Australian bryophytes vary within very wide limits, depending on whether each recorded binomial is treated as a species unless it has been formally reduced to synonymy. This treatment inflates the number of species actually present. The alternative of recognising only those species that are judged to be different on the basis of specimens examined will lead to an under-estimate but is probably much closer to reality, except in the case of groups that have not yet been studied properly. The number of species still awaiting discovery is purely conjectural but is likely to be of the order of 200–300 in each group. The lower and upper limits might be: mosses: 600 (Scott 1982a), and 1200 (Streimann & Curnow 1989); liverworts: 400 (Scott 1982a) and 600 (Scott & Bradshaw 1986). The lower figures include names treated as synonyms until proved valid, the higher treat all names as accepted until proved synonymous. The difference consists very largely of those species that are known only from the type specimen or description. Most of them were described more than 50 years ago and therefore contribute to the potential X category. In the list of bryophytes given in Appendix B, such nomenclatural species are omitted and only the few that are known with reasonable certainty to be 'sound' taxa and which meet the criteria for extinction are included in Category X. Only a few of them, that are well established cases, are given here. The rest in the list are mostly Endangered (by reason of great rarity and hence susceptibility to human activities) or, in the case of slightly more abundant species, Vulnerable or Potentially Vulnerable. Some, of course, will be not rare but rarely collected.
The basic principle underlying endangered species legislation is that every species should be able to survive and retain its evolutionary potential in the wild. To this end, the Commonwealth Endangered Species Protection Act 1992, is aimed at, inter alia, 'promoting the recovery of species and ecological communities that are endangered or vulnerable' and 'preventing other species and ecological communities from becoming endangered'. Knowledge of bryophytes and their distributions is too incomplete for a list of endangered species to be practicable but the list of species known to be rare is likely to include most of those that are at risk within the known flora. The following (Appendix B) list includes those species of bryophyte that have been recorded only once or twice and a few others that are reasonably suspected of being in need of protection, omitting a few ostensibly rare species whose rarity is reasonably believed to be illusory. There are many other pre-1950 records of supposedly new species, known only from the type collection. The taxonomy is dubious and all early records have therefore been omitted. The categories (Ramsay & Seur 1990) are those of ANZECC with the suffixes discussed above (Introduction 1.3) which can be dropped off, if so desired, without affecting the major categories. Because the great majority of very rare mosses are the discoveries of Dr Ilma Stone, who has spent many years working on the moss flora of Queensland in particular, their categories tend to reflect the number of times she has collected them. They are mostly very sporadic in their occurrence and a true estimate of the extent to which they are endangered is not feasible. The additional coding of rarity is that used in Briggs & Leigh (1989).
Most of the Xt and Et species, which are rather large categories, have been omitted from the list to save space. Further research will clearly transfer them to other categories.
3.4. Threatening processes
For bryophytes, virtually all threatening processes involve loss of habitat on either a macro or a micro scale. (1) Historically, removal of forest and conversion to agricultural land has been the greatest single process responsible for loss of bryophyte habitat. Today this threat continues but an increasingly important cause of loss is probably the change to drier soil and drier microclimate, brought about by unsuitable forest management (including habitat fragmentation, breaches of canopy cover, large coupe size and too-rapid felling rotation); (2) too frequent burning (which increases the probability of coinciding with a subsequent drought, thus diminishing recovery); and (3) alteration to flooding regimes by removal of water for irrigation (e.g. in the Wimmera) or other flood-control practices. Salination of semi-arid regions (4) also causes loss of habitat and, although it simultaneously creates new habitat of a type favourable to some of the rarer Australian bryophytes, there does not seem to be any appreciable colonisation by them. Bryophyte habitats can also be lost (5) by natural processes such as succession, windthrow, drought, natural fire, flood, erosion etc.
Most of these processes, except the natural ones, have been largely under State control, and early bryophyte collections were few in number and limited in area, so the losses of bryophytes due to them are much easier to document on a State-wide basis.
Land clearance
What has been lost by land clearance for agricultural is unknowable. The best indications perhaps come from the Richmond R. region of New South Wales where W.W. Watts collected from 1896 to 1903 (Ramsay 1980). Of the 167 binomials of mosses alone for which Watts collected the type material, a quite high proportion (c. 20%) were collected in the Richmond R. region, in forested habitats that were later cleared. Many have never been collected since, although there has been no exhaustive survey. Subsequent taxonomic revisions of some genera have shown that very many of these, perhaps as much as half, are not worthy of specific rank, but even so the permanent loss of species is likely to have been substantial.
A similar situation is found in the mallee. The remaining mallee of Victoria, South Australia and New South Wales is one of the richest bryophyte habitats, particularly in the minute and often ephemeral mosses and liverworts of soil crusts. Almost no collecting was done in these woodlands in the days before the great clearances, and the loss of species, although probably large, is unprovable. Clearance of, and damage to, mangrove woodlands are less important for bryophytes than for lichens (Chapter 2) but bryophytes do occur there and the communities have not yet been surveyed adequately or even explored (Windolf 1989).
Too-frequent burning
There is only anecdotal evidence of the effects on bryophytes of what seems to be increasingly frequent, and now often annual, burning in Cape York and other parts of Queensland, but there are very few hard data. Some mosses have apparently disappeared from localities (e.g in Tasmania) that are now burnt frequently; cause and effect may be inferred but not proved.
Alteration to flooding regimes
Substantial areas of the Wimmera drainage system and adjacent Victorian mallee are now never flooded and other areas very rarely so, apparently because of the water being tapped off from the Wimmera River for irrigation. Although no rare species is known to have suffered as a result, the comparatively great biodiversity of areas that are still flooded (such as the Stawell flats) makes it likely that there have been significant losses. Other areas of both drainage and irrigation (e.g., Bool Lagoon, S.A., and along the Murray and Murrumbidgee Rivers) are of sufficient size to make damage to cryptogam communities inevitable even in the absence of detailed records.
Salination
Although the most widespread effect of this is where non-saline soils become saline following a rise in the water table, destroying any pre-existing bryophytes, even salt-tolerant bryophytes can be killed by changes in the salinity regime. Riella halophila, growing in the Lochiel salt lake (Pink Lake), Dimboola, Vic., was inundated with saline water when the lake flooded in 1977 and, together with Funaria salsicola, was killed and has not yet returned although the water level soon receded to its previous level.
Natural causes
Treubia tasmanica was apparently lost to Victoria when a landslide in the early 1970s wiped out one of the only two known colonies. The other seems to have been lost through its roadside habitat drying out.
Riella spiculata, known only from a boggy meadow near Portland, Vic., disappeared after 1952 when the paddock dried up. It has not been seen since, although searches have been casual and intermittent. It is unknown whether the loss is permanent, but the paddock is now far from boggy. The precise locality, of course, may have been misidentified, and a wider search would be worthwhile.
3.5. Habitat conservation
The micro-habitat requirements for individual species are almost wholly unknown and the knowledge we have is based on casual observation and opinion, not on measurement. As an example, we know that Tortula papillosa and T. pagorum are mosses that grow together particularly in rain-tracks on the trunks of trees (especially non-eucalypts) although also found on e.g. vertical concrete walls. The proportion of T. pagorum in the mixture appears to increase the lower the rainfall (Scott & Stone 1976); the community is absent from dark and/or very wet forest and is commonest in open, well-lit habitats where the distinction between rain-tracks and other parts of the trunk is pronounced. At that level, the ecology is fairly well known but there are no quantitative data whatsoever on the temperature, rainfall, and light climate that delimit the habitat, or on those that apparently correlate with changing proportions of the two components. In such a prevailing state of knowledge, the prescriptions of habitats and their conservation requirements can only be in the broadest terms.
Within this limitation, the principal bryophyte habitats in Australia would appear to be:
Rainforest (tropical and temperate)
Ground species, epiphylls and epiphytes on trunks, branches and upper canopy twigs.
Semi-wet or dry temperate forests
Predominantly ground bryophytes, although Frullania, Radula and Lejeuneaceae may be abundant on favourable phorophytes such as Acacia in the wettest forests in this category.
Semi-arid (including mallee)
Soil crust bryophytes where the soil is moist in the growing season and retains sufficient moisture for growth and development to be completed. A rich, varied flora (Scott 1982a, 1982b).
Stabilised sandy soils, e.g. fixed sand dunes
The flora is comparable to 3 above, but less rich in species. Where dune scrub is sufficiently dense, there may be epiphytes.
Grasslands
Bryophytes are commonly confined to (and abundant in) moist sheltered micro-habitats, commonly in the lee of tussocks or more generally dispersed amongst grasses in moister grasslands.
Roadsides
Flora of 3, 4 and 5 above, depending on locality.
Rocks
Characteristic floras develop on limestone and granite, particularly where the rocks are sufficiently exposed to be independent of tree canopy.
Aquatic
The richest flora in this category is in rapidly flowing streams in wet forest; a much more restricted flora in slow-moving waters; and a very small but highly specialised flora, both floating and stranded on mud, in still water bodies. Marshes and bogs have a characteristic but small flora, different from any of these.
Saline
Very few, highly specialised and distinctive species survive in this habitat and are confined to it. There are very few bryophytes indeed in salt marshes (Adam 1976; Windolf 1989) and fewer still in mangrove swamps.
Mountain/alpine
The flora of rocks, rock crevices and soil above the tree line and amongst grasses in alpine/subalpine meadows is rich, varied, distinctive but under-investigated.
Based on present knowledge, several localities are particularly rich in rare and endangered species, corresponding to Churchill's concept of centres of endemism arising from islands in the Australian Tertiary sea. These are Mt Wellington in Tasmania, Mts Gower and Lidgbird on Lord Howe Is., and Mt Bellenden Ker, Thornton Peak and Mt Lewis in Queensland. There may well be more when the flora is better known.
The full effects of the bushfires that burnt out Mt Wellington in 1967 have not been recorded, but recovery of some flowering plants has been very slow, so the habitat is still not recovered to a condition where an inventory of bryophyte losses could be compiled with certainty.
3.6. Improving conservation measures
3.6.1 Impediments to conservation
Impediments affecting bryophytes in particular, rather than organisms in general, fall under three headings.
1. Unrecognisability
On the part of the general public, most mosses and liverworts are not recognised as such; and those that are, are generally not threatened. There is, therefore, no discriminating public driving force that can be specifically focused on bryophyte conservation. On the part of conservation workers and support scientists there is a lack of taxonomic expertise, a state slowly being changed by the provision of literature, tuition courses and heightened awareness; but expertise is never likely to be widespread, because of the nature of the organisms, their small size and often transient nature which impose limits to recognisability comparable to those of many invertebrates.
These are inherent problems that can probably never be solved but only, as far as possible, circumvented; for example by the efficient use of a small number of specialists.
2. Unknown ecology
The ecology of almost every species of bryophyte in Australia, and certainly every rare or endangered species, is unknown and can only be conjectured, at best, by a scrutiny of the habitat. We know nothing of their response to fire, flood, drought, or disturbance, nor to changes of light and humidity; nothing of their longevity, reproduction, colonisation, powers of recovery and spread. As with almost all plants we do not know why they die. Lack of knowledge of basic responses makes their management wildly haphazard. A possible strategy to improve this position is proposed below.
3. Subordinate role
With the probably sole exception of Sphagnum bogs and possibly also some soil crusts, bryophytes occupy a subordinate ecological role in their communities. They can therefore be conserved only by conserving the entire community. But to put it in polarised terminology, it is obviously administratively and politically difficult to conserve an otherwise worthless community for the sake of the worthwhile bryophytes within it. On the other hand, if the major botanical components of the community are worth conserving in their own right, it may be difficult for the bryophytes not to be given lower priority in any conservation plan.
In other words, bryophytes can only realistically be conserved by conservation of the entire habitat, with a concomitant awareness of their significance as components of the conserved community.
3.6.2. Possible strategy
1. As suggested above, ignorance of bryophyte taxonomy and ecology are major impediments to conservation. Realistically, the best position that can be hoped for is a general appreciation among conservation workers of the existence and ecological significance of bryophytes, a small number of bryologists capable of advising, and an affordable encouragement of research in taxonomy and especially ecology. Within this modest framework it ought to be possible, in principle, with a few years' work, to develop a set of guidelines on likely bryophyte responses to changing habitat factors and management processes; guidelines based on observation, not experiment, and hence only equivalent to best current advice, not to scientific proof. This would involve an assessment of the bryophyte components of all different kinds of community in Australia, and their likely responses to expected perturbations, on the basis of present knowledge, however flimsy that might be.
2. The lack of bryological taxonomic expertise amongst botanists, and the present inadequate information on the ecology and distribution of bryophytes, interact. If the proposal for maximising the use of a few specialists, as suggested in the paragraph above, were to be adopted, this would also make it possible to produce assessments of sites, as suggested in Section 1.6, i.e. a register of SSSIs. This would then provide a body of data on which decisions, as well informed as current knowledge would allow, could be made on bryophyte conservation.
3. Where there is inadequate knowledge of biological and ecological factors (as is the case with most species), research will be required to establish the basic data. In many cases this research need not be either in great depth or for an extended time. This sort of work could be done most cost-effectively by grants at honours student level, to provide travel costs and small items of equipment, for several months' work under an appropriate supervisor or co-supervisor. In the past, travel costs in particular have proved a major obstacle to taxonomic and ecological work, even though the sums involved are not great. This is an equivalent scheme to the Small Research Grants schemes already being run by the British and Australian Ecological Societies which have proved capable of yielding useful, productive results. Where prolonged observations are required, of course, this sort of collaboration would not be feasible and either a postgraduate studentship or a collaboration with a conservation agency would be required.
3.7. Public awareness
Possible mechanisms to establish improved understanding have been outlined in the Introduction, Sections 1.6, 1.7; in particular, completion of the three bryological volumes of the Flora of Australia, production of wall charts and posters and simple handbooks for general botanists at all levels from school to post-university. For many years an elementary book of moss and liverwort photographs and accompanying text has been in production by Mr B.A. Fuhrer and Miss D.C. Cargill at Monash University but has been held up partly by lack of funds. If this could be completed and produced at a low price, it would be an excellent way of spreading knowledge of bryophytes to a wider public.
Cards of undesignated type that could be used for Christmas cards, birthday cards or note-cards, and widely marketed, could be another way of increasing public awareness. The ABRS card of Petalophyllum preissii, painted by Celia Rosser, produced in the 1980s was a good example, but was not widely available.
References
Adam, P. (1976). The occurrence of bryophytes on British salt marshes. Journal of Bryology 9: 265–274.
Ando, M. & Matsuo, A. (1984). Applied Bryology. Advances in Bryology 2: 133–224.
Banwell, A. D. (1951). A new species of Riella from Australia. Transactions of the British Bryological Society 1: 475–478.
Bates, J. W. (1990). Interception of nutrients in wet deposition by Pseudoscleropodium purum: an experimental study of the uptake and retention of potassium and phosphorus. Lindbergia 15: 93–98.
Briggs, J. D. & Leigh, J. H. (1989). Rare or threatened Australian plants, 1988 revised edn. Australian National Parks and Wildlife Service, Canberra.
Burges, A. (1932). Notes on the mosses of New South Wales. I. Additional records and description of a new species of Buxbaumia. Proceedings of the Linnean Society of New South Wales 57: 239–244.
Catcheside, D. G. (1980). Mosses of South Australia. Government Printer, Adelaide.
Catcheside, D. G. (1987). The genus Bartramia (Bartramiaceae) in Australia. Memoirs of the New York Botanical Garden 45: 618–626.
Catcheside, D. G. & Stone, I. G. (1988). The mosses of the Northern Territory. Journal of the Adelaide Botanic Gardens 11: 1–17.
Clymo, R. S. (1963). Ion exchange in Sphagnum and its relation to bog ecology. Annals of Botany n.s. 27: 309–324.
Clymo, R. S. (1982). The Ecology of Sphagnum. pp. 229–289 in Bryophyte Ecology ed by A. J. B.Smith. Chapman & Hall, London & New York
Coley, P. D., Kursar, T. A. & Machado, J-L. (1993). Colonization of tropical rainforest leaves by epiphylls: effects of site and host plant leaf lifetime. Ecology 74: 619–623.
Cropper, S. C., Tonkinson, D. A. & Scott, G. A. M. (1991). A census of Victorian Bryophytes. 56pp. Victorian Government Publ., Melbourne.
Dalton, P. D., Seppelt, R. D., & Buchanan, A. M. (1991). An annotated checklist of Tasmanian mosses. pp. 15–32 in Aspects of Tasmanian Botany – A tribute to Winifred Curtis ed by M. R. Banks et al. Royal Society of Tasmania, Hobart.
Engel, J. J. (1987). Miscellanea Hepaticologica Paleotropica. I. Memoirs of the New York Botanical Garden 45: 533–534.
Engel, J. J. (1991). Studies in Tasmanian Hepaticae. I. The genus Adelanthus. pp. 7–11 in Aspects of Tasmanian Botany – A tribute to Winifred Curtis ed by M. R. Banks et al. Royal Society of Tasmania, Hobart.
Geissler, P., Tan, B. & Hallingbäck, T. (1993). Towards a World Red List of Bryophytes. (Unpub. ?).
Gradstein, S. R. (1992). Threatened bryophytes of the neotropical rainforest: a status report. Tropical Bryology 6: 83–93.
Grolle, R. (1962). Goebelobryum eine neues marsupiale Lebermoosgattung. Journal of the Hattori Botanical Laboratory 25: 135–144.
Grolle, R. (1982). Ubersicht der Lejeuneaceae in Tasmanien. Wissenschaftliche Zeitschrift der Friedrich-Schiller-Universität Jena/Thüringen. Mathematisch-naturwissenschaftliche Reiche 31: 207–222.
Grolle, R. (1987). Die Lebermoosgattung Lepidolaena in Queensland. Haussknechtia (Jena) 3: 65–69.
Hattori, S. (1979). A revision of the Australasian species of the genus Frullania, Hepaticae. I. Journal of the Hattori Botanical Laboratory 45: 323–363.
Hattori, S. (1983). A revision of the Australasian species of Frullania Hepaticae III. Journal of the Hattori Botanical Laboratory 54: 133–184.
Hattori, S. (1984). Marie Hicks' Frullania collection made in Northern Queensland. Cryptogamie – Bryologie et Lichénologie 5: 177–190.
Hattori, S. (1987a). New or little known species of Frullania (Frullaniaceae) from Queensland and New South Wales, Australia. Memoirs of the New York Botanical Garden 45: 544–555.
Hattori, S. (1987b). The Frullania flora of Lord Howe Island. Bryologist 90: 365–370.
Hattori, S. (1988). Frullania collections made by Dr. Barbara M. Thiers in Queensland and New South Wales, Australia. Beihefte zur Nova Hedwigia 90: 147–158.
Hedenäs, L. & Soderström, L. (1992). Endangered Bryophytes in Europe-cause and conservation. Biological Conservation 59, special issue: 85–270.
Hewson, H. J. (1970a). The family Aneuraceae in Australia and New Guinea. I. The genus Aneura. Proceedings of the Linnean Society of New South Wales 94: 173–193.
Hewson, H. J. (1970b). The family Aneuraceae in Australia and New Guinea. II. The genus Riccardia. Proceedings of the Linnean Society of New South Wales 95: 60–121.
Hewson, H. J. (1982). Vandiemeniaceae: a new family in the Metzgeriales. Journal of the Hattori Botanical Laboratory 52: 163–165.
Hicks, M. L. (1991). Scapania in Queensland, Australia. Journal of the Hattori Botanical Laboratory 69: 129–132.
Hodgetts, N. G. (1992). Guidelines for selection of biological SSSIs: non-vascular plants. Joint Nature Conservation Committee, UK.
Inoue, H. (1986). Notes on the Plagiochilaceae XIII. Review of the genus Plagiochila (Dum.) Dum. in mainland Australia. Journal of the Hattori Botanical Laboratory 60: 357–378.
Kantvilas, G. & Jarman, S. L. (1991). Lichens and bryophytes of the Tasmanian World Heritage Area. 1. Mount Sprent. pp. 149–162 in Aspects of Tasmanian Botany – A tribute to Winifred Curtis ed by M. R. Banks et al. Royal Society of Tasmania, Hobart.
Lai, M-J. (1992). Pseudospiridentopsis horrida (Musci, Trachypodaceae), a neglected moss genus and species from Australia. Journal of Japanese Botany 67: 44–47.
Lewinsky, J. (1990). Zygodon Hook. & Tayl. in Australasia: a taxonomic revision including SEM – studies of peristomes. Lindbergia 15: 109–139.
Moore, C. J. & Scott, G. A. M. (1979). The ecology of mosses on a sand dune in Victoria, Australia. Journal of Bryology 10: 291–311.
Norris, D. H. & Montalvo, A. M. (1981). The rediscovery of Rhabdodontium buftonii (Broth. & Geh. in Broth.) Broth. Bryologist 84: 85–88.
Norris, D. H. & Robinson, H. (1981). Stoneobryum, a new genus of Orthotrichaceae from South Africa and Queensland. Bryologist 84: 95–99.
Ochyra, R. (1986). Touwia laticosta, a remarkable new genus and species of moss from Queensland, Australia. Journal of Bryology 14: 103–108.
Ochyra, R. (1990). Streimannia turgida, a new genus and species of Brachytheciaceae from Australia. Journal of Bryology 16: 223–229.
Ochyra, R. (1991). Bryostreimannia nom. nov. replaces Streimannia Ochyra, hom. illeg. Journal of Bryology 16: 643–644.
Olarinmoye, S. O. (1974). Ecology of epiphyllous liverworts: growth in three natural habitats in Western Nigeria. Journal of Bryology 8: 275–289.
Ramsay, H. P. (1980). Contributions of Rev. W. W. Watts F.L.S. to Australian botany. Taxon 29: 455–469.
Ramsay, H. P. (1984a). Census of New South Wales mosses. Telopea 2: 455–534.
Ramsay, H. P. (1984b). The mosses of Lord Howe Island. Telopea 2: 549–558.
Ramsay, H. P. & Seur, J. (1990). Type specimens of bryophytes in Australian herbaria. pp. 247–252 in History of systematic botany in Australasia ed by P. S. Short. Australian Systematic Botany Society Inc., South Yarra.
Ramsay, H. P. & Streimann, H. (1984). Mosses and their distribution in the Australian Capital Territory. Telopea 2: 559–574.
Ramsay, H. P., Coveny, R. G., Brown, E. A. & Brooks, A. K. (1993). Bryophytes of the Royal Botanic Gardens, Government House and the Domain, Sydney. Cunninghamia 3: 215–229.
Ratkowsky, D. A. (1987). Checklist of Tasmanian liverworts. Papers & Proceedings of the Royal Society of Tasmania 121: 153–158.
Ratkowsky, D. A. & Ratkowsky, A. V. (1982). The bryophytes of the Mt Wellington Range, Tasmania. Papers & Proceedings of the Royal Society of Tasmania 116: 97–115.
Schuster, R. M. (1971). Studies on Cephaloziellaceae. Nova Hedwigia 22: 121–266c.
Schuster, R. M. & Engel, J. J. (1974). A monograph of the genus Pseudocephalozia (Hepaticae). Journal of the Hattori Botanical Laboratory 38: 665–701.
Schuster, R. M. & Engel, J. J. (1985). Austral Hepaticae V(2). Temperate and subantarctic Schistochilaceae of Australia. Journal of the Hattori Botanical Laboratory 58: 255–539.
Scott, G. A. M. (1982a). Bryofloristics in Australasia. Nova Hedwigia 71: 483–494.
Scott, G. A. M. (1982b). Desert Bryophytes. pp. 105–122 in Bryophyte Ecology ed by A. J. E. Smith. Chapman & Hall: London & New York.
Scott, G. A. M. (1985). Southern Australian Liverworts. Australian Government Publishing Service, Canberra.
Scott, G. A. M. & Bradshaw, J. A. (1986). Australian liverworts (Hepaticae): annotated list of binomials and check-list of published species with bibliography. Brunonia 8: 1–171.
Scott, G. A. M. & Pike, D. C. (1984). New species of Fossombronia from Australia. Journal of the Hattori Botanical Laboratory 56: 339–349.
Scott, G. A. M. & Pike, D. C. (1987). Studies on Fossombronia in Australia II. Fourteen more new species. Journal of the Hattori Botanical Laboratory 62: 367–386.
Scott, G. A. M. & Pike, D. C. (1988). A new species of Fossombronia in Australia. Beihefte zur Nova Hedwigia 90: 159–162.
Scott, G. A. M. & Stone, I. G. (1976). The Mosses of Southern Australia. Academic Press, London.
Stone, I. G. (1973). A new species of Brachydontium from Australia. Journal of Bryology 7: 343–351.
Stone, I. G. (1975). Trachycarpidium in Queensland, Australia. Muelleria 3: 122–129.
Stone, I. G. (1976). A remarkable new moss from Queensland, Australia – Viridivellus pulchellum new genus and species (new family Viridivelleraceae) Journal of Bryology 9: 21–31.
Stone, I. G. (1977). Bruchia queenslandica, a new moss from tropical Australia. Journal of Bryology 9: 509–518.
Stone, I. G. (1982a). Some new and noteworthy records of mosses mostly from Queensland, Australia. Austrobaileya 1: 511–520.
Stone, I. G. (1982b). Erpodium australiense (Erpodiaceae), a new species of moss from Australia. Journal of Bryology 12: 191–197.
Stone, I. G. (1983). Buxbaumia in Australia, including a new species B. thorsborneae. Journal of Bryology 12: 199–208.
Stone, I. G. (1984a). Notes on four species of moss which are common to South Africa and Australia. Journal of Bryology 13: 129–130.
Stone, I. G. (1984b). Uleobryum curtisii sp. nov. (Pottiaceae) from Queensland, Australia. Journal of Bryology 13: 19–24.
Stone, I. G. (1985a). Archidium thalliferum sp. nov. with a persistent cushion-shaped protonema unique in Musci. Journal of Bryology 13: 345–352.
Stone, I. G. (1985b). New records of mosses in Australia. Journal of Bryology 13: 475–478.
Stone, I. G. (1985c). Calymperastrum, a new genus of Pottiaceae. Journal of Bryology 14: 315–318.
Stone, I. G. (1985d). A comparison of the species of Fissidens subgenus Aneuron in Australia. Journal of Bryology 14: 319–325.
Stone, I. G. (1987). Tristichella dimorpha (Sematophyllaceae), a new species from Queensland. Journal of Bryology 14: 691–699.
Stone, I. G. (1989). Revision of Phascum and Acaulon in Australia. Journal of Bryology 15: 745–777.
Stone, I. G. (1990). Nomenclatural changes and new moss record in Australia: including a description of the protonema of Calomnion. Journal of Bryology 16: 261–273.
Stoneburner, A., Wyatt, R., Catcheside, D. G. & Stone, I. G. (1993). Census of the mosses of Western Australia. Bryologist 96: 86–101.
Streimann, H. & Curnow, J. (1989). Catalogue of mosses of Australia and its external territories. Australian Government Printer, Canberra.
Streimann, H. & Touw, A. (1981). New records for some Australian mosses. Journal of the Hattori Botanical Laboratory 49: 261–271.
Takakai, N. & Ando, H. (1975). Rare species and their conservation: Bryophyta. pp. 105–108 in Studies in conservation of natural terrestrial ecosystems in Japan ed by M. Numata, K. Yoshioka & M. Kato. Japanese International Biosphere Program Synthesis Vol. 8. University of Tokyo Press, Tokyo.
Taylor, J. (1954). A new Australian hepatic. Kew Bulletin 9 (1): 45–47.
Thiers, B. M. (1986). Austrolejeunea bidentata, a new species of Lejeunaceae Subfamily Tuyamaelloideae from Australia. Bryologist. 88: 350–352.
Thiers, B. M. (1987a). Lepidolejeunea queenslandica (Lejeuneaceae subfamily Lejeunioideae): A new species from Australia. Memoirs of the New York Botanical Garden 45: 556–560.
Thiers, B. M. (1987b). A preliminary account of Colura (Hepaticae, Lejeuneaceae) in Australia. Brittonia 39: 175–179.
Thiers, B. M. (1988). The Australian species of Cololejeunea. Beih. Nova Hedwigia 90: 113–146.
Thiers, B. M. & Gradstein, S. R. (1989). Lejeuneaceae (Hepaticae) of Australia I. Subfamily Ptychanthoideae. Memoirs of the New York Botanical Garden 52: 1–81.
Vitt, D. H. & Ramsay, H. P. (1985). The Macromitrium complex in Australasia (Orthotrichaceae: Bryopsida). Part 1. Taxonomy and phylogenetic relationships. Journal of the Hattori Botanical Laboratory 59: 325–451.
Whinam, J., Eberhard, S., Kirkpatrick, J. & Moscal, T. (1989). Ecology and conservation of Tasmanian Sphagnum peatlands. Tasmanian Conservation Trust.
Willis, J. H. (1950). The chequered story of two Tasmanian mosses. Victorian Naturalist 67: 30–35.
Windolf, J. (1987). Annotated check list of Queensland Hepaticae. Austrobaileya 2: 380–400.
Windolf, J. (1989). Bryophytes in a sub-tropical mangrove community. Austrobaileya 3: 103–107.
Wyatt, R. & Stoneburner, A. (1989). Pleurophascum occidentale: a new moss from Western Australia. Bryologist 92: 299–301.
Yamada, K. (1987). A preliminary study of the genus Radula from Queensland, Australia. Journal of the Hattori Botanical Laboratory 62: 191–200.
Yamaguchi, T., Seppelt, R. D., Iwatsuki, Z. & Buchanan, A. M. (1990). Sphagnum (sect. Buchanania) leucobryoides sect. et sp. nov. from Tasmania. Journal of Bryology 16: 45–54.
