1948-

2018

Ray Specht: A Retrospective

Editor David Doley

The Royal Society of Queensland

6 December 2018 Revised 16 January 2019 The International Biological Program The International Geosphere – Biosphere Program

By Raymond Louis Specht Emeritus Professor of Botany, The University of Queensland Email < [email protected] >

Editor: David Doley Centre for Mined Land Rehabilitation, The University of Queensland Email < [email protected]>

CONTENTS Summary Introduction: Understanding Ecosystem Structure and Function

1. Ecosystem Productivity 1.1. Structure and productivity of Australian plant communities 1.2. Foliage Projective Cover 1.3. Vegetation cover and condition in Queensland

2. Ecosystem Production Processes 2.1. Solar radiation interception by foliage canopies 2.2. Annual photosynthesis and evapo-transpiration from plant communities 2.3. Rainfall interception by foliage canopies and resultant runoff 2.4. Evaporative Coefficient 2.5. Climate, vegetation height and species diversity 2.6. Seasonal growth rhythms of foliage in overstorey and understorey 2.7. Closed vegetation communities 2.8. Regeneration of arid zone plant communities after overgrazing during drought 2.9. Biochemical processes in Australian plant communities 2.9.1 Phosphorus biochemistry of vegetation on nutrient-poor soil 2.9.2 Phosphorus toxicity in vegetation on nutrient-poor soil 2.9.3 C4 Photosynthetic Pathway 2.9.4 Sodium as a micro-nutrient for C4 plants 2.9.5 Stomatal resistance, measured by the Carbon Isotope Ratio 2.9.6 Leaf Specific Weight (dry weight per unit area) and Leaf Area 2.9.7 Chlorophyll a/b ratio 2.9.8 Nitrate reductase enzyme activity 2.9.9 Resurrection chloroplasts in the leaves of arid zone grasses 2.9.10 Calcium carbonate toxicity 2.9.11 Altitudinal cline of Eucalyptus pauciflora (Snow Gum)

3. Ecosystem Conservation 3.1 Classification of Australian plant communities 3.2 Conservation of Australian plant communities 3.3 Species richness of vascular plants and resident vertebrates 3.3.1 Alpha Biodiversity (Species Richness) of vascular plants and resident vertebrates 3.3.2 Beta Biodiversity (Species Richness along ecological gradients) 3.3.3 Gamma Biodiversity (Species Richness per region)

1 3.4 Seed dispersal 3.5 Species evolution in seepage of uraniferous radiation 3.6 Palaeobotanical studies in Australia 3.6.1 Climate and soils 3.6.2 Late Cretaceous palynology 3.6.3 Mid Tertiary palynology 3.6.4 Quaternary palynology and vegetation 3.7 Rare and endangered Australian plants 3.8 Selection of conservation reserves 3.9 Conservation by cultivation

4. Syntheses 4.1 Ethno-botany 4.2 American-Australian Scientific Research Expedition to Arnhem Land (1948) 4.3 Ecosystems of the world 4.4 Ecological biogeography of Australia 4.5 Mediterranean-type ecosystems 4.6 Australian plant communities. Dynamics of structure, growth and biodiversity

5. Education 5.1 Teaching material for Queensland tertiary students in botany 5.2 Biological Science curriculum studies – ‘The Web of Life’ 5.3 University of Queensland’s 75th celebrations

6. Community services 6.1 Childhood asthma — associated with seasonal growth of eucalypt foliage 6.2 The mining industry 6.2.1 Bauxite mining, Weipa 6.2.2 Fluoride emission 6.2.3 Sulphur dioxide emission from copper refiners 6.2.4 Serpentinite, a nickel-bearing deposit 6.2.5 Mine-reclamation 6.3 Soft-rot in electricity poles after the 1974 Brisbane flood 6.4 Growing trees on the Brisbane tuff 6.5 Mt Coot-tha Botanic Gardens

7. Conclusions

8. Publications of Raymond Louis Specht

2 Summary

As part of the International Biological Program, Ray Specht performed a central role in Australia-wide studies of the Dynamics of Structure, Growth and Biodiversity of Plant Communities (with resident vertebrates — small mammals, birds, snakes, reptiles and amphibia), together with their Conservation. A lifetime of work resulted in many new findings in each of the IBP topic areas.

Systematic description of plant community structure and its response to environmental conditions was a fundamental concern. It became clear that the fundamental connection between community structure and environment was the influence of the atmospheric conditions during the season of shoot growth. The basically positive effect of increasing temperature on shoot extension growth can be offset by the frictional, and evaporative aerodynamic fluxes generated as air flows over and through a plant community. Increasing fluxes both desiccate and abrade new foliage growth at the edges of foliage clusters, crown units or the whole crowns of large plants and the crowns of individuals or groups of small plants in a vegetation community. The distribution of this growth restriction within a plant community results in a balance between the Foliage Projective Covers in the overstorey (FPCo) and understorey (FPCu), so that Σ (FPCo + FPCu) is a constant for all plant communities within a particular climatic area — along all edaphic gradients and during post-fire successions.

The photosynthetic energy fixed annually by the combined Foliage Projective Covers determines the species-richness of vascular plants and resident vertebrates (small mammals, birds, snakes, lizards and Amphibia) in a mature ecosystem.

Species-richness of plants slowly declines during a post-fire succession, but is re-activated by fire.

An increase in ambient temperature of 2oC is predicted to increase the atmospheric aerodynamic fluxes that affect shoot development throughout a plant community, and thus influence the Foliage Projective Cover. As a result, changes in plant community structure may be predicted to occur as follows:

Open-forest —> woodland —> tall shrubland —> low shrubland

Tall open-forest —> open-forest —> Savannen-Wald —> Savannen-Gebusch —> Savanne

Heide-Wald —> Sklerophyllen-Wald —> Sklerophyllen-Gebusch —> Heide

Rainforests, monsoon forests, dry scrubs will survive in well-watered habitats

This degradation is associated with decreasing rainfall-interception by the canopy, thus, increasing run-off from the ecosystem.

Plant communities were evaluated with the help of the TWINSPAN computer package. The conservation and survival of the 397 TWINSPAN Floristic Groups — containing the majority of Australian plant species with associated consumers and decomposers — was regarded as essential, not only in conservation reserves but in response to the community-physiological stresses of Global Warming. In 1995, only 37% of these TWINSPAN Groups were conserved, ranging from 70% of rainforests to only 11% of savanna Groups.

3 Based on the inferred influence of Global Warming in the Mid-Tertiary, rainfall is predicted to increase over the next 50 years in the northern part of Australia, resulting in an increase in vertical growth of foliage shoots and, thus, the annual photosynthetic production per hectare. However, increasing temperature will increase aerodynamic fluxes that reduce Foliage Projective Cover of the canopy, reduce canopy interception of rainfall, and increasedrun-off.

As the continent of Australia is drifting northwards at about 66 mm per year, monsoonal troughs may bring heavy rainfalls as far south as Sydney (latitude 34oS). This change in temperature and rainfall distributions will ensure the regeneration of mulga (Acacia aneura) and other Arid Zone species in the south of their range.

Rainfall in southern Australia is predicted to decrease with global warming, forcing the Mediterranean- type climate systems southward and threatening the survival of many producers, consumers and decomposers in each of the TWINSPAN Floristic Groups. However, evolution of new plants may occur in well-watered sites and, thus, increase the Gamma Biodiversity of south-west Western Australia.

With Global Warming, a rise in sea level will affect all estuaries around Australia, especially the Black Soil Plains of the Top End that will be submerged by the high tidal-range — the habitat of many birds will be reduced.

4 INTRODUCTION: Understanding Ecosystem Structure and Function

Seven years after UNESCO identified research into arid zones as a priority activity, CSIRO and the UNESCO Arid Zone Research Program held a symposium on Climatology and Microclimatology in Canberra in 1956 that brought together physicists, soil scientists and plant and animal biologists from around the world (UNESCO, 1958). Australian scientists (including Specht 1958a) made many seminal contributions to the understanding of the state and movement of water in the soil and atmosphere, and consequences for both plants and animals of arid zones. As a result, the CSIRO Division of Meteorological Physics formulated an international program to integrate micro-climatology and plant ecology to measure evapotranspiration, net photosynthesis (during the day) and respiration (at night) continuously throughout the day (and year) — a program that they promoted through the World Meteorological Organisation.

During 1964, while Ray Specht held a Royal Society–Nuffield Foundation Fellowship in the Agriculture Department of Oxford University, Prof. Geoffrey Blackman — on behalf of the Royal Society — and Professor R.N. ‘Bob’ Robertson, the Biological Secretary of the Australian Academy of Science, nominated him to serve on three Planning Committees of the International Biological Program: Section PT (Productivity), Section PP (Production Processes), and Section CT (Conservation) of Terrestrial Ecosystems. During the next 18 months, Planning Committees met in Paris, France (PT), Wageningen, Netherlands (PP) and Monk’s Wood, England (CT).

The Australian Academy of Science subsequently appointed Specht as Convenor of these three sections of the International Biological Program (IBP). He was then encouraged to move to subtropical Queensland to promote research relevant to the three IBP Sections on Terrestrial Ecosystems —the structure, growth and biodiversity of terrestrial ecosystems and their conservation throughout Australia. This work formed the core of his interests for a further 50 years.

1. ECOSYSTEM PRODUCTIVITY

1.1 Structure and Productivity of Australian Plant Communities IBP Section PT, on the Productivity of Terrestrial Plant Communities, aimed to measure the biomass of plant communities, above and below ground, and assess the annual nutrient-cycling within these two strata, searching for the ecological mechanisms proposed by Watt (1947) and elaborated for woodlands by Ovington (1962).

The study of the biomass and nutrient-status of roots was very difficult in tall temperate eucalypt forests growing on hard clay soils (Ashton 1976a; Attiwill 1966, 1968. 1979, 1980). In contrast, the team studying Dark Island heathland (Specht et al. 1958) was easily able to study root systems in the deep sandy soil under this vegetation. Similarly, Walt Westman and Rod Rogers were able to study the root-system of a subtropical eucalypt forest in a pit created purposely by sand-miners on North Stradbroke Island, Queensland (Westman & Rogers 1977a, 1977b).

While further studies were inhibited by the extent of site disturbance required and by the physical and financial demands of adequate sample collection, this Australian work conformed to the generalisation

5 that the ratio of biomass in roots to tops increases progressively in plant communities from the humid to the arid climatic zone (Rodin 1972).

References Ashton, D. H. (1976). The development of even-aged stands of Eucalyptus regnans F. Muell. in Central Victoria. Australian Journal of Botany 24: 397-414. Attiwill, P. M. (1966). A method of estimating crown weight in Eucalyptus, and some implications of relationships between crown weight and stem diameter. Ecology 47: 795-804. Attiwill, P. M. (1968). Determination of forest biomass. Forest Science 14: 13-15. Attiwill, P. M. (1979). Nutrient cycling in a Eucalyptus obliqua (L’Hérit.) forest. III. Growth, biomass and net primary production. Australian Journal of Botany 27: 439-458. Attiwill, P. M. (1980). Nutrient cycling in a Eucalyptus obliqua (L’Hérit.) forest. IV. Nutrient uptake and nutrient return. Australian Journal of Botany 28: 199-222. Carter, J. O. (1996). The relationship between Foliage Projective Cover and Basal Area of overstorey trees in over 150 timbered stands from the arid to the perhumid zone in Queensland. Queensland Department of Primary Industries. Resource Management Institute, Brisbane. Ovington, J. D. (1962). Quantitative ecology and the woodland ecosystem concept. Advances in Ecological Research 1: 103-192. Rodin, L. (Ed.) (1972). Eco-physiological Foundation of Ecosystem Productivity in the Arid Zone. Publishing House ‘Nauka, Leningrad. Specht, R. L., Rayson Patricia & Jackman, Margaret E. (1958). Dark Island heath (Ninety-Mile Plain, South Australia). 6. Pyric succession: Changes in composition, coverage, dry weight, and mineral nutrient status. Australian Journal of Botany 6: 59-88. Watt, A. S. (1947). Pattern and process in the plant community. Journal of Ecology 35: 1-22. Westman, W. E. & Rogers, R. W. (1977a). Biomass and structure of a subtropical eucalypt forest, North Stradbroke Island, Queensland. Australian Journal of Botany 25: 171-191.

1.2. Foliage Projective Cover A critical element of both the structural and functional description of a plant community is the amount of incident solar radiation that can be intercepted by defined components of that community (Monteith 1965). While light interception by plant canopies is ideally measured by radiometers, this is feasible only for experimental situations, and a more rapidly applicable estimate of plant canopy leaf area was needed. A workable solution was the estimation of the foliage projective cover, which is the percentage of sky obscured by foliage (not stems) above a point of observation. Crosswire sighting tubes enabled precise assessments of the Foliage Projective Covers of both overstorey (FPCo) and understorey (FPCu) strata of plant communities throughout Australia (Specht 1970; Howard 1970).

The Foliage Projective Covers (horizontal foliage covers per hectare) of the overstorey and understorey strata were found to be in equilibrium (Specht and Morgan 1981). Several consequences of this relationship were:

 The Foliage Projective Cover of the understorey (FPCu) is determined by the overstorey (FPCo) so that (FPCo + FPCu) remained constant throughout post-fire succession (Specht et al. 1958).  The same relationship between overstorey FPCo and understorey FPCu was observed along edaphic gradients where shallower soils retained less soil water than in deeper soils (Specht et al. 2005, 2006).   (FPCo + FPCu) remained constant as half the original 30 Myoporum platycarpum trees that were on Q100 on Koonamore Vegetation Reserve in 1926, died by 1962 (Hall et al. 1964).  FPCo + FPCu) was found to remain constant in Thinning Experiments on mesquite in Arizona (Parker & Martin 1952) and on mulga at Charleville. (Burrows & Beale 1969).

6   (FPCo + FPCu) was found to be a constant at every climatic location, correlated with the Evaporative Coefficient (Specht 1972a).

Plant community structure is influenced by aerodynamic fluxes (frictional, thermal, evaporative) in the atmosphere as wind flows over and through a plant community (Marshall 1971). These fluxes abrade the growth of new foliage shoots at the top and sides of overstorey and understorey plants throughout every plant community, leading to more compact crown shapes in dry as compared with humid environments (Specht & Yates 1990: Specht et al. 1991).

Paradoxically, seasonal shoot growth of overstorey plants in southern and south-eastern Australia occurs during the driest season of the year, whereas shoot growth in understorey plants favours the wet seasons (Specht & Rayson, 1957; Specht & Brower 1975; Rogers & Westman1981; Westman 1978). Walt Westman (1978) considered that the overstorey eucalypt and understorey heathy strata were of two distinct origins. However, David Yates and Ray Specht showed that, as internode-lengths decreased in species in the gradient from overstorey to the ground, more solar radiation was trapped momentarily in the axils of leaves, thus inducing foliage growth earlier (Specht & Yates 1990; Specht et al. 1991).

References Burrows, W. H. & Beale, I. F. (1969). Structure and association in the mulga (Acacia aneura) lands of south-western Queensland. Australian Journal of Botany 17: 539-552. Hall, E., Anne A., Specht, R. L. Eardley, Constance M. (1964). Regeneration of the vegetation on Koonamore Vegetation Reserve, 1926-62. Australian Journal of Botany. 12: 205-264. Howard, J. A. (1970). Stereoscopic profiling and the photogrammetric description of woody vegetation. Australian Geographer 11: 359-372. Marshall, J. K. (1971). Drag measurements in roughness arrays of varying density and distribution. Agricultural Meteorology 8: 269-292. Monteith, J. L. (1965). Light distribution and photosynthesis in field crops. Annals of Botany 29(1): 17-37. Parker, R. K. & Martin, S. C. (1952). The Mesquite Problem on Southern Arizona Ranges. United States Department of Agriculture, Circular No. 908. Rogers, R. W. & Westman, W. E. (1981). Growth rhythms and productivity of a coastal subtropical eucalypt forest. Australian Journal of Ecology 6: 85-98. Specht, R. L. (1970). Vegetation. Pp. 44-67. In: The Australian Environment. Fourth Edition. (Ed. G. W. Leeper) CSIRO & Melbourne University Press, Melbourne. Specht, R. L. (1972). Water use by perennial, evergreen plant communities in Australia and Papua New Guinea. Australian Journal of Botany 20: 273-299. Specht, R. L. & Brouwer, Yvonne M. (1975). Seasonal shoot growth of Eucalyptus spp. in the Brisbane area of Queensland (with notes on shoot growth and litter fall in other areas of Australia). Australian Journal of Botany 23: 459-474. Specht, R. L. & Morgan, D. G. (1981). The balance between the foliage projective covers of overstorey and understorey strata in Australian vegetation. Australian Journal of Ecology 6: 193-202. Specht, R. L. & Rayson, Patricia (1957). Dark Island heath (Ninety-Mile Plain, South Australia). 1. Definition of the ecosystem. Australian Journal of Botany 5: 52-85. Specht, R. L. & Yates, D. J. (1990). Climatic control of structure and phenology of foliage shoots in dicotyledonous overstorey and understorey strata of subtropical plant communities in eastern Australia. Acta Oecologia 11: 215-233. Specht, R. L., Rayson, Patricia & Jackman, Margaret E. (1958). Dark Island heath (Ninety-Mile Plain, South Australia). 6. Pyric succession: Changes in composition, coverage, dry weight, and mineral nutrient status. Australian Journal of Botany 6: 59-88. Specht, R. L., Yates, D. J., Sommerville, Jacqui E .M. and Moll, E. J. (1991). Foliage structure and shoot growth in heathlands in the mediterranean-type climate of southern Australia and South Africa. Ecologia Mediterranea 16 : 195-207. Specht, R. L., Batianoff, G. N. & Reeves, R. D. (2005). The vegetation continuum on the serpentinite soil catena in the Rockhampton – Marlborough area. Pp. 23-40. In: Marlborough Serpentine, Landscape Management and Conservation. (Eds Anne Exelby, K. Wormington and A. Melzer) Centre for Environmental Management, Central Queensland University, Rockhampton. Specht, R. L., Batianoff, G. N. & Reeves, R. D. (2006). Vegetation structure and biodiversity along the eucalypt forest to rainforest continuum on the serpentinite soil catena in a subhumid area of Central Queensland, Australia. Austral Ecology 31: 394-407.

7 Westman, W. E. (1978). Evidence for the distinct evolutionary histories of canopy and the understorey in the Eucalyptus forest-heath alliance of Australia. Journal of Biogeography 5: 365-376.

1.3. Vegetation cover and condition in Queensland The assessment of vegetation cover and condition are essential components of any evaluation of ecosystem productivity. Traditional ground-based survey methods had produced broad-scale vegetation maps, but the periodic assessment of vegetation condition and potential ecosystem productivity had been beyond the reach of researchers in the 1950s and 1960s. A new approach became possible during the course of the IBP Ecosystem Productivity project.

Vegetation and land surfaces reflect the various wavebands of visible and infrared solar radiation in ways that are related to their contents of water or chlorophyll per unit land area. Tucker (1979) developed the Normalised Difference Vegetation Index (NDVI) as a function of the reflectances of visible red radiation

NDVI = (infrared – red) / (infrared + red) While this form of analysis was almost impossible from ground-based or conventional aerial platforms, it became relatively straightforward after the commissioning of the satellites, including the USA National Oceanic and Atmospheric Administration (NOAA) satellites (e.g., Tarpley et al. 1984; Justice et al. 1985). The NDVI, calculated for each pixel of a satellite image, was closely correlated with the total green biomass visible in each pixel (Tucker 1979). NDVI is also related closely to leaf area index up to a value of about 2, and with fractional vegetation cover (or foliage projective cover) (Carlson & Ripley 1997). These identities allowed rapid and repeated large-scale assessment of vegetation that could detect both seasonal effects and land use changes (e.g., Nelson & Holben 1986).

The relationship between FPCo and NDVI was assessed in Queensland from satellite imagery by John Carter, Tim Danaher and others of the Statewide Land-cover and Tree Study (SLATS) (with Ray Specht, who had recently completed a ‘Conservation Atlas of Plant Communities in Australia’, on the Advisory Committee). ‘The SLATS Reference Group was established to provide feedback from a wide range of stakeholders and to assist with the communication of results to industry and the wider community’ (DERM 2010, P. 5). This extensive survey enabled tree-clearing to be assessed throughout the State of Queensland from 1989 to 2010 (DERM 2010).

Over the same period, research staff of the Queensland Herbarium assessed the clearing of remnant vegetation from 1995 to 2010 (Accad et al. 2008). Between 1988 and 2009, 8,460 x 1,000 hectares of forest-woodland vegetation were cleared throughout Queensland. Since 1995, 57% of a total of 5,140 x 1,000 hectares of cleared land was remnant vegetation. Land clearing in Queensland has contributed a significant proportion of Queensland’s and Australia’s total greenhouse gas emissions (Henry et al. 2002; DERM 2010).

8 The degradation of Australia’s rangelands during drought and their recovery after rain was followed by Franks et al. (2002) and McKeon et al. (2004). The leaves of many inland grasses appear dead on the onset of drought, but their chloroplasts resurrect as soon as rains fall (Doley & Trivett 1974; Mittelheuser 1977).

References Accad, A., Neldner, V. J., Wilson, B. A. & Niehus, R. E. (2008). Remnant vegetation in Queensland: Analysis of remnant vegetation 1997-2005, including regional ecosystem information. Queensland Herbarium, Environmental Protection Agency, Brisbane. Armston, J. D., Denham, R. J., Danaher, T. J., Scarth, P. F. & Moffet, T. (2000). Prediction and validation of foliage projective cover from Landsat-5TM and Landsat-7TM ETM+ imagery. Journal of Applied Remote Sensing 3: 033540. Carlson, T. N. & Ripley, D. A. (1997). On the relation between NDVI, fractional vegetation cover, and leaf area index. Remote sensing of Environment, 62(3): 241-252. Carter, J. O. (1996). The relationship between Foliage Projective Cover and Basal Area of overstorey trees in over 150 timbered stands from the arid to the perhumid zone in Queensland. Queensland Department of Primary Industries. Resource Management Institute, Brisbane. Department of Environment and Resource Management (DERM) (2010). Land cover change in Queensland 2008-09: a Statewide Landcover and Trees Study (SLATS) Report, 2011. Queensland Department of Environment and Resources Management, Brisbane. Doley, D. & Trivett, N. B. A. (1974). Effects of low water potentials on transpiration and photosynthesis in Mitchell Grass (Astrebla lappacea). Australian Journal of Plant Physiology 1: 539-550. Franks, A. J., Playford, Julia & Shapcott, Alison (Eds) (2002). Landscape Health of Queensland. The Royal Society of Queensland. Henry, Beverley K., Danaher, T. J., McKeon, G. M. & Burrows, W. H. (2002). A review of the potential role of greenhouse gas abatement in native vegetation management in Queensland’s rangelands. Rangeland Journal 24: 112-132. Justice, C. O., Townshend, J. R. G., Holben, B. N. & Tucker, E .C., 1985. Analysis of the phenology of global vegetation using meteorological satellite data. International Journal of Remote Sensing, 6(8), pp.1271-1318. McKeon, G., Hall, W., Henry, Beverley, Stone, G. & Watson, I. (2004). Pasture Degradation and Recovery in Australia’s Rangelamds. Learning from History. Queensland Department of Natural Resources, Mines and Energy, Brisbane, Queensland. Mittelheuser, Cathryn J. (1977). Rapid ultrastructural recovery of water-stressed leaf tissue. Zeitschrift fur Pflanzenphysiologie 82: 458-461. Nelson, R. & Holben, B., 1986. Identifying deforestation in Brazil using multiresolution satellite data. International Journal of Remote Sensing, 7(3), pp.429-448. Specht, R. L., Specht, Alison, Whelan, M. B. & Hegarty, Elwyn E. (1995). Conservation Atlas of Plant Communities in Australia. Southern Cross University, Lismore, New South Wales. Tarpley, J. D., Schneider, S. R. and Money, R .L., 1984. Global vegetation indices from the NOAA-7 meteorological satellite. Journal of Climate and Applied Meteorology 23(3): 491-494. Tucker, C. J. (1979). Red and photographic infrared linear combinations for monitoring vegetation. Remote Sensing of the Environment 8: 127-150.

2. ECOSYSTEM PRODUCTION PROCESSES

2.1. Solar radiation interception by foliage canopies

On the first day of Ray Specht’s B.Sc. Hons. project to survey the distribution of eucalypt species in the Mount Lofty Ranges (Specht and Perry 1948), he studied the eucalypts along the ridge at the end of Green Hill Road, Adelaide. This north-south facing ridge ran from the Adelaide Plain (with rainfall about 25 inches per annum) towards Mt Lofty (with rainfall about 35 inches per annum). At the lower rainfall, Eucalyptus leucoxylon occupied the north-facing slope while E. viminalis occupied the shaded south-facing slope. As rainfall increased, E. viminalis displaced E. leucoxylon on the sunny north-facing slope.

“The influence of topography on the incidence of solar radiation on foliage canopies was therefore ingrained in my mind”.

In preparation for the first ANZAAS Conference to be held in Adelaide after the Second World War, Prof. Joe G. Wood asked Ray to make a balsa model showing the distribution of eucalypt species in his study

9 area of the Adelaide Hills. “My model was a great success with interstate visitors”. Prof. John S. Turner of the University of Melbourne encouraged Trevor Clifford to study the ecology of eucalypts in part of the Dandenong Range and illustrate their distribution on a model (Clifford 1954). The distribution of eucalypt species on the slopes of the isolated volcanic cone of Mount Piper demonstrated the effect of solar radiation in all points of the compass (Ashton 1976a).

During the 1950s, Ray Specht and his students studied the mineral nutrition of heathy vegetation that had survived for 50-100 million years on soils very low in phosphates since the Late Cretaceous (Specht and Rayson 1957; Specht et al. 1958). To the west of the relatively flat sandplain where we established our Dark Island heath experiments (Specht 1963), a clump of Eucalyptus baxteri trees nestled in the eastern side of a barchan dune. M.Sc. student, Patricia Rayson, was encouraged to map the vegetation on a grid- pattern across the barchan dune (Rayson 1957). To learn more about the micro-meteorology of the site, Ray Specht approached Bruce Mason of the South Australian office of the Bureau of Meteorology. The Bureau lent them ten standard rain-gauges to examine the monthly rainfall at sites on a west to east transect across the barchan dune; they found that rainfall falling in westerly winds was swept upwards to be deposited in the easterly-facing arc of the barchan, thus enabling E. baxteri trees to survive (Rayson 1957).

As well, Bruce Mason asked the CSIRO Division of Building Research to lend the group their model ‘Solarscope’ to enable them to plot mean monthly incident solar radiation falling on the various sections of a model of the barchan dune. The ‘Solarscope’ had been developed by CSIRO staff to plot the pattern of solar rays that would fall on architectural models in different latitudes and geographical orientations (Phillips 1948).

The ‘Solarscope’ became a teaching-tool for ecology classes in the University of Adelaide during the 1950s, then in the University of Queensland in the late 1960s.

When the University of Queensland installed a PDP10 computer, lecturer, David Yates, developed a computer program to simulate the physical ‘Solarscope’. This program enabled an estimation of monthly solar radiation falling on localities on every month at different latitudes and geographical orientation and slope. Long-term monthly cloud records could adjust the solar radiation falling on foliage canopies. This program was used extensively for both teaching and research in the Botany Department.

In tropical Australia, the sun shines from the north during summer and from the south during winter. The growth of lichens, mosses, liverworts and most ferns, which grow on the ground, die due to overheating (Rogers and Stevens 1981).

David Yates down-loaded the program for Latitude 60oS to enable Ray Specht, Mary Dettmann and David Jarzen to postulate the structure of vegetation that occurred in south-eastern Australia when it was attached to Antarctica at the end of the Late Cretaceous, with a subtropical climate (Specht et al. 1992).

References Ashton, D. H. (1976). The vegetation of Mount Piper, central Victoria: A study of a continuum. Journal of Ecology 64: 463-483. Clifford, H. T. (1954). On the distribution of the species of Eucalyptus in the region of the Dandenong Range, Victoria. Proceedings of the Royal Society of Victoria 65: 30-55.

10 Phillips, R. O. (1948). Sunshine and Shade in Australasia. 1st ed. (Duplicated Document No 23). Sydney: Commonwealth Experimental Building Station. Rayson, Patricia (1957). Dark Island heath (Ninety-Mile Plain, South Australia). II. The effects of micro-topography on climates, soils and vegetation. Australian Journal of Botany 5: 85-102. Rogers, R. W. and Stevens, G. Nell (1981). Lichens. Pp. 591-603. In: Ecological Biogeography of Australia. (Ed. A. Keast). Junk, The Hague. Specht, R. L. (1963). Dark Island heath (Ninety-Mile Plain, South Australia). 7. The effect of fertilizers on composition and growth, 1950-1960. Australian Journal of Botany 11: 67-94. Specht, R. L. and Perry, R. A. (1948). Plant ecology of part of the Mount Lofty Ranges. Transactions of the Royal Society of South Australia 72: 91-132. Specht, R. L., Rayson, Patricia and Jackman, Margaret E. (1958). Dark Island heath (Ninety-Mile Plain, South Australia). 6. Pyric succession. Changes in composition, coverage, dry weight and mineral nutrient status. Australian Journal of Botany 6: 155-176. Specht, R. L., Dettmann, Mary E. and Jarzen, D. M. (1992). Community associations and structure in the Late Cretaceous vegetation of south-east Australasia and Antarctica. Palaeogeography, Palaeoclimatology, Palaeoecology 94: 283-309.

2.2. Annual Photosynthesis and Evapotranspiration from Plant Communities When Specht was nominated as convenor of the International Biological Program Section on Physiological Processes in Terrestrial Ecosystems for the International Biological Program, he was astounded to learn that the Committee comprised micro-meteorologists, a computer modeler, plant physiologists, crop- physiologists, as well as ecologists dealing with forests, grasslands and natural plant communities. It was his job to formulate an integrated program that would satisfy these diverse members. David Angus from the CSIRO Division of Meteorological Physics, Tom Neales, a crop-physiologist from the University of Melbourne, and Specht, a plant ecologist from the University of Adelaide, produced a program (Specht 1966, 1967) that satisfied the IBP steering committee, with little alteration.

By the time that Specht moved to Queensland, in 1967, the CSIRO Division of Meteorological Physics had developed a meteorological mast of micro-instruments to measure temperature, water vapour and carbon dioxide concentrations continuously over a plant community throughout the day. The data were recorded in the field by a computer, large for the time with a random-access memory of 8 kb, housed in a caravan and power by a portable generator. All this equipment was very expensive, but a grant was obtained from the newly instituted Australian Research Grant Committee (ARGC).

A micro-meteorologist, a biophysicist and a plant biochemist formed the interdisciplinary team of scientists appointed to study different ecosystem components, as well as to develop teaching courses in the Botany Department of the University of Queensland. The assembly of such a diverse team of scientists, and implementation of the inevitably divergent research and teaching programs, resulted in a loss of focus on the ecosystem process objectives of the IBP project.

Then, it was learnt that the theoretical assumptions underlying the processing of data obtained by instruments on the CSIRO meteorological mast applied only to smooth-surfaced plant communities such as grasslands with an upwind extent of more than two km. A grant was obtained to study a uniform sward of Mitchell grass near Charleville in south-western Queensland. This project was abandoned on the advice of Champ Tanner, a visiting micro-meteorologist from the University of Wisconsin, U.S.A., who doubted whether the micro-meteorological measurements made in such an erratic rainfall climate would be useful.

11 An area of short grassland on one edge of Archerfield Aerodrome, Brisbane, was then selected and a lysimeter installed to compare direct evapotranspiration measurements with the records obtained from a meteorological mast. However, this project was abandoned when David Angus returned to Victoria.

Aspects of the work planned for Charleville continued at the plant level, particularly on the rapid recovery of photosynthesis and transpiration in curly Mitchell grass (Astrebla lappacea) following the imposition and relief of severe water deficits (Doley and Trivett 1974), which was explained by the maintenance of integrity and rapid recovery of chloroplast structure and function through a desiccation cycle (Mittelheuser 1977). Mitchell grass also indicated ecological tolerance by adapting quickly and extensively to changes in growth regime temperature (Doley and Yates 1976).

Parallel studies of tissue water relations (Connor and Tunstall 1968), tree photosynthesis (Connor et al. 1971), tree water balance and plot-level hydrology (Tunstall and Connor 1975) had been conducted in brigalow (Acacia harpophylla) forest near Meandarra, western Queensland, an environment renowned for variation between flooded soils and severe water deficits. This work indicated the limits to seasonal and annual photosynthetic capacity of brigalow forest that underpin estimates of ecosystem dry matter productivity.

References Connor, D.J. and Tunstall, B.R. (1968). Tissue water relations for brigalow and mulga. Australian Journal of Botany, 163, 487-490. Connor, D.J., Tunstall, B.R. and Driessche, R. (1971). Analysis of photosynthetic response in a brigalow forest. Photosynthetica. 5, 210-217 Doley, D. & Trivett, N. B. A. (1974). Effects of low water potentials on transpiration and photosynthesis in Mitchell Grass (Astrebla lappacea). Australian Journal of Plant Physiology 1: 539-550. Doley, D. and Yates, D.J. (1976). Gas exchange of Mitchell grass (Astrebla lappacea (Lindl.) Domin.) in relation to irradiance, carbon dioxide supply, leaf temperature and temperature history. Australian Journal of Plant Physiology, 3, 471-487. Mittelheuser, C.J. (1977). Rapid ultrastructural recovery of water stressed leaf tissue. Zeitschrift für Pflanzenphysiologie, 82, 458-461. Specht, R. L. (1958). Micro-environment (soil) of a natural plant community. In: Climatology and Microclimatology, Proceedings of the Canberra Symposium. Arid Zone Research, Vol. XI, 152-155. Paris, France. Specht, R. L. (1966). The International Biological Program (I.B.P.), Section PCT: Productivity, production processes and conservation of terrestrial communities. Australian Journal of Science 28: 377-380. Specht, R. L. (1967). The photosynthesis of plant communities in relation to structure, physiology and environment. Photosynthetica 1: 132-134. Specht, R. L. (2011). Development of ecosystem research. International Scholarly Research Network (ISRN Ecology) Vol. 2011, Article ID 897578, 20 pages. Tunstall, B.R. and Connor, D.J. (1975). Internal water balance of brigalow (Acacia harpophylla F. Muell.) under natural conditions. Australian Journal of Plant Physiology, 2, 489-499. UNESCO (1958). Climatology and Microclimatology, Proceedings of the Canberra Symposium. Arid Zone Research Vol. XI. Paris, France. Westman, W. E. and Rogers, R. W. (1977b). Nutrient stocks in a subtropical eucalypt forest, North Stradbroke Island, Queensland. Australian Journal of Ecology 2: 447-460.

2.3. Rainfall interception by foliage canopies and resultant runoff

The importance of rainfall interception by tree canopies and its transfer to the soil through stemflow was demonstrated clearly in Mediterranean heath vegetation in south Australia (Specht 1957) and in mulga communities of arid central Australia (Slatyer 1965) and semi-arid western Queensland (Pressland 1973, 1976). In these environments, the concentration of water falling on tree crowns within the root zones of trees and shrubs was considered to be important for their survival through the dry season.

12 Subtropical rainforests of southern Queensland are restricted to increasingly higher altitudes with increasing distance from the coast. This phenomenon is associated with the occurrence of cloud and the interception of cloud water by a rainforest canopy and its transfer to the ground through fog drip. While stemflow associated with rain events was substantial in this forest, throughfall was often recorded in the absence of gauge rainfall, sometimes adding one-third of the total annual gauge rainfall (Hutley et al. 1997). In addition, the maintenance of wet leaf surfaces overnight is sufficient to eliminate leaf water deficits in Sloanea woollsii, a subtropical rainforest canopy tree species (Yates and Hutley 1995). The addition of fog drip to the soil water supply beyond the meteorological rainfall total is considered to be an important contributor to the water balance and survival of forests with high FPCo in relatively low rainfall environments.

References Hutley, L.B., Doley, D., Yates, D. J. and Boonsaner, A. (1997). Water balance of an Australian subtropical rainforest at altitude: the ecological and physiological significance of intercepted cloud and fog. Australian Journal of Botany, 45, 311•329. Pressland, A.J., 1973. Rainfall partitioning by an arid woodland (Acacia aneura F. Muell.) in south-western Queensland. Australian Journal of Botany, 21(2), pp.235-245. Pressland, A.J., 1976. Soil moisture redistribution as affected by throughfall and stemflow in an arid zone shrub community. Australian Journal of Botany, 24(5), pp.641-649. Slatyer, R. O. (1965). Measurements of precipitation interception by an arid zone plant community (Acacia aneura F. Muell.). Arid Zone Research 25: 181-192. Slatyer, R. O. (1967). Plant-Water Relationships. Academic Press, New York. Specht, R. L. (1957). Dark Island heath (Ninety-Mile Plain, South Australia). 4. Soil moisture patterns produced by rainfall interception and stem-flow. Australian Journal of Botany 5: 137-150. Specht, R. L. and Specht, Alison (1999 Hardback; 2002 Paperback). Australian Plant Communities. Dynamics of Structure, Growth and Biodiversity. Oxford University Press, Melbourne. Yates, D.J. and Hutley, L.B., (1995). Foliar uptake of water by wet leaves of Sloanea woollsii, an Australian subtropical rainforest tree. Australian Journal of Botany, 43: 157-167.

2.4. Evaporative Coefficient The soil moisture patterns that were produced by stem-flow in the overstorey plants in Dark Island heath — Banksia ornata and Xanthorrhoea australis — were followed at monthly intervals for three years (Specht 1957a,b). It was clear that soil water accumulated during the wettest period of the year was used during the dry summer season by this evergreen vegetation. The monthly ratios of Moisture index, or Actual to Potential evapotranspiration, when plotted against available soil water in each month, formed a linear regression. The slope of this linear regression was termed the Evaporative Coefficient. Post-graduate students, Helene Martin, Bailey Carrodus and Richard Jones, were encouraged to determine the Evaporative Coefficients, respectively, of the evergreen eucalypt forest vegetation at Inglewood in the Adelaide Hills (Martin and Specht 1962), saltbush and bluebush shrublands on Koonamore Station in north-eastern South Australia (Carrodus and Specht 1965), and heathland at Frankston, near Melbourne (Specht and Jones 1971). The linear regression lines of the Evaporative Coefficients formed an array from the Arid to the Humid Climatic Zones.

For a hydrologically neutral site, where annual rainfall equals annual actual evapotranspiration, there is no net gain or loss of water from the site by lateral or vertical flow, and there is no month during which the root-extractable soil water store is completely exhausted. The annual integral of the monthly evaporative coefficients is equal to the ratio of annual precipitation, P, to annual potential evapotranspiration, Ep, (which can usually be approximated by free water evaporation, Ew). This identity (P/E) means that estimates of the evaporative coefficients of evergreen plant communities

13 could be made at every long-term climate station in Australia and then plotted on a map of Australia (Specht 1972a,b). This identity also allowed mature perennial plant communities with different structural and compositional attributes to be assessed in terms of the site water balance. Importantly, this expression allowed for hydrological variation (maximum water storage, runoff and drainage) to be included in the calculation of site water balance and equilibrium vegetation characteristics.

References Carrodus, B. B. and Specht, R. L. (1965). Factors affecting the relative distribution of Atriplex vesicaria and Kochia sedifolia (Chenopodiaceae) in the arid zone of South Australia. Australian Journal of Botany 13: 419-433. Martin, Helene A. and Specht, R. L. (1962). Are mesic communities less drought resistant? A study on moisture relationships in dry sclerophyll forest at Inglewood, South Australia. Australian Journal of Botany 10: 106-118. Specht, R. L. (1957a). Dark Island heath (Ninety-Mile Plain, South Australia). 4. Soil moisture patterns produced by rainfall interception and stem-flow. Australian Journal of Botany 5: 137-150. Specht, R. L. (1957b). Dark Island heath (Ninety-Mile Plain, South Australia). 5. The water relationships in heath vegetation and pastures on the Makin Sand. Australian Journal of Botany 5: 151-172. Specht, R. L. (1972a). Water use by perennial, evergreen plant communities in Australia and Papua New Guinea. Australian Journal of Botany. 20: 273-299. Specht, R. L. (1972b). Water use by plant communities in the arid zone of Australia. pp. 48-53 In Eco-physiological Foundation of Ecosystem Productivity in the Arid Zone. Publishing House "Nauka", Leningrad. Specht, R. L. and Jones, R. (1971). A comparison of the water use by heath vegetation at Frankston, Victoria and Dark Island Soak, South Australia. Australian Journal of Botany 19: 311-326.

2.5. Climate, vegetation height and species diversity Attributes of plant communities, including vegetation height and species diversity, are associated with climate in ways that can be explained by energy and mass exchange relationships. Shoot extension at the edges of foliage canopies is influenced by frictional, thermal and evaporative fluxes (Marshall 1971) that may either promote or restrict apical expansion at the precise location of the shoot extremity (Specht and Yates 1990). In addition, species with relatively slender stems and small shoot apices that are not robust, protected by scales or subtending foliage are subject to abrasion from adjacent canopies, producing a phenomenon recognised as crown shyness. These aerodynamic fluxes at the edges of canopies determine the Foliage Projective Cover in the plant community. In contrast, the growth of vertical shoots is determined by available soil water, available soil nutrients, and temperature (Specht and Specht 1989a). Under an optimal environment for plant growth, the integral of Leaf Area Index (total area of all leaves per unit ground surface area) determines the total amount of solar energy intercepted over a year and, therefore, Alpha (local) species diversity (Whittaker 1972; Specht and Specht 1989b). A gradient in Leaf Area Indices was found from Humid to Arid Climatic Zones; from deep soils to shallow soils; from rich soils to nutrient-poor soils; from tropical to temperate climates; from sea-level to alpine altitudes in Australia (Specht and Specht 1989a, 1999).

References Marshall, J. K. (1971). Drag measurements in roughness arrays of varying density and distribution. Agricultural Meteorology 8: 269-292. Specht, R. L. and Specht, Alison (1989a). Canopy structure in Eucalyptus-dominated communities in Australia along climatic gradients. Acta Oecologia. Oecologia Plantarum 10: 191-202. Specht, R. L. and Specht, Alison (1989b). Species richness of overstorey strata in Australian plant communities — The influence of overstorey growth rates. Australian Journal of Botany 37: 321-336. Specht, R. L. and Specht, Alison (1999, 2002). ‘Australian Plant Communities. Dynamics of Structure, Growth and Biodiversity’ (Oxford University Press, Melbourne).

14 Specht, R. L. and Yates, D. J. (1990). Climatic control of structure and phenology of foliage shoots in dicotyledonous overstorey and understorey strata of subtropical plant communities in eastern Australia. Acta Oecologia, Oecologia Plantarum 11: 215-233. Whittaker, R. H. (1972). Evolution and measurement of species diversity. Taxon. 21: 213-251

2.6. Seasonal Growth Rhythms of Foliage in Overstorey and Understorey While Ray Specht and students were studying the rooting patterns of species in the bone-dry soil under Dark Island heath during December 1950 — the beginning of summer in the Mediterranean- type climate of South Australia — they noticed that new shoots were appearing on the bushes of the dominant Banksia ornata. Every month over the next two years, the number of leaves on selected shoots and their characteristics were measured. At the same time, records were made of new shoots and flowers on all component species in the heath. Overstorey species produced new leaves during late spring through summer; understorey species only grew during spring (Specht and Rayson 1957).

Many years later, it was recalled that the leaves of understorey species were more closely packed, with shorter internode lengths, than in the overstorey species. Using micro-instrumentation in the laboratory, Specht and Yates (1990) found that incident solar radiation was absorbed by the densely- packed leaves, raising the temperature of the enclosed shoot apex, thus inducing shoot-growth earlier than in overstorey species that had long internodes, exposed apices and more effective re-radiation of energy. In effect, all component species in the heath had the same separation between spring and summer growth rhythms (Specht et al. 1975, 1991).

Helene Martin undertook a soil moisture study of the dry sclerophyll forest extending from a humid to a sub-humid climate along an east-west ridge at Inglewood in the Mount Lofty Ranges. The growth rhythm of the eucalypt trees occurred during the summer season when soil moisture rapidly declined, faster in the humid climate than in the subhumid climate (Martin and Specht 1962). This conundrum encouraged Russell Sinclair to study the water potential and stomatal conductance of leaves of three species of Eucalyptus in the Adelaide Hills throughout the year (Sinclair 1980).

The leaves in the canopy of eucalypt forests in South Australia and Victoria (Specht and Specht 1989) — later in south-west Western Australia, thanks to Joe Havel — all had the summer growth rhythm in a Mediterranean-type climate. Specht et al. (1992) and Specht and Specht (1995, 1999) hypothesised that these plant communities had retained this ‘subtropical’ growth rhythm since the continent of Australia had been attached to Antarctica (lat. 60oS) in the Late Cretaceous with a palaeo-temperature of 15-20oC (Shackleton and Kennett 1975) and drifted northwards at an average rate of 66 mm per annum (Wellman and McDougall 1974). The Mediterranean-type climate was produced by the path of the westerly winds blowing across the Southern Ocean moving northward towards the Equator in summer-time (Specht et al. 1991).

The seasonal shoot-growth of the eucalypt forest in south-eastern Queensland retains the ‘subtropical’ rhythm of southern Australia — but, now, with growth in autumn and spring, not in the hotter summer

15 or the cooler winter seasons (Specht and Brouwer 1975). Further north, the foliage growth-rhythm changes to ‘tropical’, with new foliage formed during summer.

The foliage rhythms of overstorey species of Acacia and understorey shrubs in the Arid Zone of South Australia appear to be similar to that of the heathy open-forests of the Mount Lofty Ranges (Lange and Sparrow 1992; Maconochie 1973, 1975; Maconochie and Lange 1970; Sinclair 1980).

References Lange, R. T. and Sparrow, A. D. (1992). Growth rates of western myall (Acacia papyrocarpa Benth.) during its main phase of canopy spreading. Australian Journal of Ecology 17: 315-327. Maconochie, J. R. (1973). Leaf and shoot growth of Acacia kempeana F. Muell. and selected arid zone species. Tropical Grasslands 7: 49-55. Maconochie, J. R. (1975). Shoot and foliage production of five shrub species of Acacia and Hakea in a dry sclerophyll forest. Transaction of the Royal Society of South Australia 99: 177-181. Maconochie, J. R. and Lange, R. T. (1970). Canopy dynamics of trees and shrubs with particular reference to the arid- zone topfeed species. Transaction of the Royal Society of South Australia 94: 243-248. Martin, Helene A. and Specht, R. L. (1962). Are mesic communities less drought resistant? A study of moisture relationships in dry sclerophyll forest at Inglewood, South Australia. Australian Journal of Botany 10: 106-118. Shackleton, N. J. and Kennett, J. P. (1975). Palaeo-temperature history of the Cenozoic and the initiation of Antarctic glaciation: Oxygen and carbon isotope analyses in DSDP sites 277, 279, 281. Pp. 743-755. In: Initial Reports of the Deep Sea Drilling Project, No. 29. United States Government Printing Office, Washington, U.S.A. Sinclair, R. (1980). Water potential and stomatal conductance of three Eucalyptus species in the Mount Lofty Ranges, South Australia: Responses to summer drought. Australian Journal of Botany 28: 499-510. Specht, R. L. (1988). Geosphere-Biosphere interaction in terrestrial ecosystems. Pp. 169-176. In: Global Change. (Ed. K. D. Cole). Australian Academy of Science, Canberra. Specht, R. L. and Brouwer, Yvonne M. (1975). Seasonal shoot growth of Eucalyptus spp. in the Brisbane area of Queensland (with notes on shoot growth and litter fall in other areas of Australia). Australian Journal of Botany 23: 459-474. Specht, R. L. and Rayson, Patricia (1957). Dark Island heath (Ninety-Mile Plain, South Australia). 1. Definition of the ecosystem. Australian Journal of Botany 5: 52-85. Specht, R. L. and Specht, Alison (1989). Canopy structure in Eucalyptus -dominated communities in Australia along climatic gradients. Acta Oecologia, Oecologia Plantarum 10: 191-213. Specht, R. L. and Specht, Alison (1995). Global warming: Predicted effects on structure and species richness of Mediterranean ecosystems in southern Australia. Pp. 215-237. In: Time Scales of Biological Responses to Water Constraints: The Case of Mediterranean Biota. (Eds J. Roy, J. Aronson and F. di Castri) SPB Academic Publishing, Amsterdam. Specht, R. L. and Specht, Alison (1999). Australian Plant Communities. Dynamics of Structure, Growth and Biodiversity. Oxford University Press, Melbourne. Specht, R. L. and Yates, D. J. (1990). Climatic control of structure and phenology of foliage shoots in dicotyledonous overstorey and understorey strata of subtropical plant communities in eastern Australia. Acta Oecologia, Oecologia Plantarum 11: 215-233. Specht, R. L., Brouwer, Yvonne M. and Derrick, E. H. (1975). Seasonal waves of asthma: A possible botanical cause. International Journal of Biometeorology 19: 28-36. Specht, R. L., Yates, D. J., Sommerville, Jacqui E. M. and Moll, E. J. (1991). Foliage structure and shoot growth in heathlands in the mediterranean-type climate of southern Australia and South Africa. Ecologia Mediterranea 16 : 195-207. Specht, R. L., Dettmann, Mary E. and Jarzen, D. M. (1992). Community associations and structure in the Late Cretaceous vegetation of southeast Australia and Antarctica. Palaeogeography, Palaeoclimatology, Palaeoecolgy 94: 283- 309. Wellman, P. and McDougall, I. (1974). Cainozoic igneous activity in eastern Australia. Tectonophysics 23: 49-65.

2.7 Closed vegetation communities If the Moisture Index, the ratio of actual to potential evaporation (Ea/Eo) closely approaches unity or the ratio of precipitation to evaporation (P/E) exceeds unity during the season of foliage growth, a closed vegetation community may develop (Specht 1972a). In Per-Humid climates, a community develops in which the total FPC is close to or even greater than 1.0 (Specht1983). The structure of the community varies with both climate and soil nutrient status. On soils with adequate nutrient availability, closed forests (rainforests) occur in the tropics (Lavarack and Stanton 1977) and

16 subtropics (Hegarty 1990, 1991; Specht, A. 1988; Stubbs 1996), while Nothofagus and conifer- dominated forests occur in temperate Australia. (Howard 1973 a, b, c).

In humid climates, wherever summer Ea/Eo equals 1.0 and average annual Ea/Eo exceeds about 0.55 (Specht 1972a), a monsoon rainforest develops in the Northern Territory (Russell-Smith 1991), and a dry scrub in subtropical Queensland (Forster et al. 1991; Specht 2007).

On nutrient-poor soils at the tip of Cape York Peninsula, a Heidewald (heathy closed-forest) develops (Lavarack and Stanton 1977), while a Kwongan (tall closed-heathland) develops on nutrient-poor soils in south-western Australia (Heddle 1980; George et al. 1979). Tall closed-heathlands are also found at the well-watered fringe of open-heathlands on nutrient-poor sands in south-eastern Australia (Specht 1979).

In the months after the closed-community had been formed when soil-water levels were optimum, fluctuations in rainfall will cause daily variations in stomatal apertures that, in turn, affect transpiration and carbon dioxide intake for photosynthesis (Doley 1981; Doley et al. 1987, 1988). Late dry-season rain events may not affect soil water storage, but they can contribute to the temporary relief of water deficits in the crowns of rainforest canopy trees.

References Doley, D. (1981). Tropical and subtropical forests and woodlands, Pp. 209-323. In: Water Deficits and Plant Growth. Vol. 4. (Ed. T. T. Kozlowski) Academic Press, New York. Doley, D., Yates, D. J. and Unwin, G. L. (1987). Photosynthesis in an Australian rainforest tree, Argyrodendron peralatum, during the rapid development and relief of water deficits in the dry season. Oecologia 74: 441- 449. Doley, D., Unwin, G. L. and Yates, D. J. (1988). Spatial and temporal distribution of photosynthesis and transpiration by single leaves in a rainforest tree, Argyrodendron peralatum. Australian Journal of Plant Physiology 15: 317-326. Forster, P. J., Bostock, P. D., Bird, L. H. and Bean, A. R. (1991). Vine-forest Plant Atlas for South-East Queensland. Queensland Herbarium, Queensland Government. George, A. S., Hopkins, A. J. M. and Marchant, N. G. (1979). The heathlands of Western Australia. Pp. 211-230. In: Ecosystems of the World. Vol. 9A. Heathlands and Related Shrublands. Descriptive Studies. (Ed. R. L. Specht) Elsevier, Amsterdam. Heddle, Elizabeth M. (1980). Effects of changes in soil moisture on the native vegetation of the northern Swan Coastal Plain, Western Australia. West Australian Forests Department Bulletin No. 92, 51 pp. Hegarty, Elwyn E. (1990). Leaf life-span and leafing phenology of lianes and associated trees during a rainforest succession. Journal of Ecology 78: 300-312. Hegarty, Elwyn E. (1991). Patterns of productivity by lianes and tree components of an Australian rainforest. Journal of Tropical Ecology 7: 201-214. Howard, Truda M. (1974a). Studies of the ecology of Nothofagus cunninghamii Oerst. I. Natural regeneration on the Mt Donna Buang massif, Victoria. Australian Journal of Botany 21: 67-78. Howard, Truda M. (1974b). Studies of the ecology of Nothofagus cunninghamii Oerst. II. Phenology. Australian Journal of Botany 21:79-92. Howard, Truda M. (1974c). Studies of the ecology of Nothofagus cunninghamii Oerst. IIII. Two limiting factors: Light intensity and water stress. Australian Journal of Botany 21: 93-102. Lavarack, P. S. and Stanton, J. P. (1977). Vegetation of the Jardine River Catchment and adjacent coastal areas. Proceedings of the Royal Society of Queensland 88: 39-48. Russell-Smith, J. (1991). Classification, species richness and environmental relations of monsoon rainforest in northern Australia. Journal of Vegetation Science 2: 259-278. Slatyer, R. O. (1978). Altitudinal variation in the photosynthetic characteristics of snow gum, Eucalyptus pauciflora Sieb, ex Spreng. VII. Relationship between gradients of field temperature and photosynthetic temperature optima in the Snowy Mountains area. Australian Journal of Botany 26: 111-121. Specht, Alison (1988). Big Scrub Conservation Strategy. Vol. 2. Resource Material. (Ed. Planners North Pty Ltd) Pp. 2.1-2.72. New South Wales National Parks and Wildlife Service, Sydney.

17 Specht, R. L. (1972). Water use by perennial, evergreen plant communities in Australia and Papua New Guinea. Australian Journal of Botany 20: 273-299. Specht, R. L. (1979). The sclerophyllous (heath) vegetation of Australia: The eastern and central States. Pp. 125- 210. In: Ecosystems of the World. Vol. 9A. Heathlands and Related Shrublands. Descriptive Studies. (Ed. R. L. Specht). Elsevier, Amsterdam. Specht, R. L. (1983). Foliage projective covers of overstorey and understorey strata of mature vegetation in Australia. Australian Journal of Ecology 8: 433-439. Specht, R. L. (2007). Species richness of rainforest stands on non-serpentinite and serpentinite substrates in the Rockhampton – Marlborough area of Central Queensland. Proceedings of the Royal Society of Queensland 113: 17-35. Stubbs, B. J. (1996). ‘A Question of Competing Values. Forest and Timber Conservation in New South Wales, 1838- 1996.’ Ph. D. Thesis, Southern Cross University, Lismore, N.S.W.

2.8. Regeneration of Arid Zone Plant Communities after Overgrazing during Drought The Federation Drought at the beginning of the 20th Century resulted in overgrazing of the native grass pastures and shrubs in the Arid Zone rangelands of inland South Australia (Osborne et al. 1935). Early in the 1920s, the owners of Koonamore Station in north-eastern South Australia gave an area of land in the most overgrazed part of one of their paddocks to the Botany Department of the University of Adelaide to monitor the regeneration of the vegetation.

Koonamore Vegetation Reserve was enclosed by a ‘rabbit-proof fence’ and permanent quadrats, line transects and photo-points were established on representative plant communities on the Reserve. Beginning in 1926, vegetation on these sites was recorded quarterly, but later annually (Wood 1936; Hall et al. 1964). These records have continued into the 21st Century — thanks to staff of the Botany Department in the University of Adelaide (Sinclair 2004, 2005).

Provided a reasonable substrate was available, seeds of saltbush (Atriplex spp.) regenerated. No regeneration of bluebush (Maireana sedifolia), that grows on harder soils, has been recorded (Carrodus and Specht 1965). The regeneration of seedlings of the False Sandalwood (Myoporum platycarpum) and Mulga (Acacia aneura) depends on monsoonal troughs to reach the area during successive seasons to secure flowering, seed maturity, germination and seedling establishment— a very rare event so far south.

Half of the 30 Myoporum trees on Quadrat 100 had died by 1960. As their canopy cover decreased, more solar radiation enabled the foliage cover of saltbush (Atriplex vesicaria) to increase linearly (Specht 1972a).

References Carrodus, B. B. & Specht, R. L. (1965). Factors affecting the relative distribution of Atriplex vesicaria and Kochia sedifolia (Chenopodiaceae) in the arid zone of South Australia. Australian Journal of Botany 13: 419-433. Hall, E. Anne A., Specht, R. L. and Eardley, Constance M. (1964). Regeneration of the vegetation on the Koonamore Vegetation Reserve, 1926 to 1962. Australian Journal of Botany 12: 205-264. Osborn, T. G. B., Wood, J. G. and Paltridge, T. B. (1935). On the climate and vegetation of the Koonamore Vegetation Reserve to 1931. Proceedings of the Linnean Society of New South Wales 50: 392-427. Sinclair, R. (2004). Persistence of dead trees and fallen timber in the Arid Zone: 76 years of data from the T.G.B. Osborn Vegetation Reserve, Koonamore, South Australia. Rangeland Journal 26: 111-122. Sinclair, R. (2005). Long-term changes in vegetation, gradual and episodic on the T.G.B. Osborn Vegetation Reserve, Koonamore, South Australia (1926-2002). Australian Journal of Botany 53: 283-296. Specht, R. L. (1972). The Vegetation of South Australia. Government Printer, Adelaide. Wood, J. G. (1936). Regeneration of the vegetation on the Koonamore Vegetation Reserve, 1926 to 1935. Transactions of the Royal Society of South Australia 60: 96-111.

18

2.9. Biochemical Processes in Australian Plant Communities As a result of the IBP Ecosystem Production Processes initiative taken up in the Botany Department, University of Queensland, a series of detailed investigations developed under the leadership of staff and students of that department and others in which Ray Specht had worked and studied. This compilation represents a brief account of the scope of these studies. 2.9.1. Phosphorus biochemistry of vegetation on nutrient-poor soil: (with Richard Groves, David Jeffrey, Wendy Haines, George Coleman). - Ortho-phosphates are released from decomposing litter under heath or eucalypt forest canopies during the moist winter-spring period in the Mediterranean-type climate of southern Australia. The soluble phosphates are taken up by and stored as poly-phosphate granules in the specialised rootlets (proteoid, cyperoid, restioid, mycorrhizal) and associated rhizosphere organisms of species adapted to these low-phosphate soils (Ashford et al. 1975; Coleman and Specht 1981). When foliage shoot apices are activated at the end of spring, poly-phosphates are hydrolysed to ortho- phosphates that are translocated to the new foliage during the dry summer season (Pate et al. 1998). 2.9.2. Phosphorus toxicity in vegetation on nutrient-poor soil: (with Peter Ozanne, CSIRO Western Australia): — An increase of ortho-phosphate in the soil increases the growth of heath plants — greatest in short-lived species; least in long-lived species. This leads to earlier death of short-lived species (Ozanne and Specht 1981). Seedling regeneration of heath plants in gaps may be inhibited by phosphorus-toxicity, leaving the site for wind- blown exotics (Specht and Specht 1999a, pp. 341-342). At Wilson’s Promontory, Victoria, coastal Leptospermum laevigatum may invade these gaps that are fertilised by ash from rubbish-fires (Burrell 1981). 2.9.3. C4 Photosynthetic Pathway: (Hal Hatch of CSR David North Research Laboratory and Botany Department, University of Queensland): — Discovered the C4 photosynthetic pathway that enabled tropical grasses and Arid Zone saltbush (Atriplex species) to survive in high-temperature and water-limited environments (Hatch and Slack 1966). Plant species in the cooler and more humid climates of Australia possess the C3 photosynthetic pathway, as do most of the plants in the cooler parts of the Northern Hemisphere. 2.9.4. Sodium as a micro-nutrient for C4 species: (Peter Brownell, Adelaide University and the Townsville Campus of the University of Queensland): — Both Arid Zone saltbush (Atriplex vesicaria) and tropical grasses (common with Africa) possessing the C4 photosynthetic pathway require sodium as a micronutrient (Brownell 1965; Brownell and Crossland 1972). Cyclic salt (sea-spray) from the ocean is blown far inland, decreasing in concentration with distance. High rates of salt deposition on vegetation near the coast have a negative effect on leaf cellular functioning, and consequently photosynthesis, plant growth and species- richness of C3 rainforests in north-eastern New South Wales, but their effects may persist well inland (Rich 1996; Specht and Specht 1999a, p.306).

19 2.9.5. Stomatal resistance, measured by the Carbon Isotope Ratio (Graham Farquhar of Australian National University, who had studied biophysics in the Botany Department, University of Queensland): The processes of gas transfer between the atmosphere and functional sites in leaves were studied and modelled for species distributed along the gradient from per-humid to arid climates (Farquhar et al. 1982a,b). 2.9.6. Leaf Specific Weight (dry weight per unit area) and Leaf Area were explored along the gradient from per-humid to arid climates by Ray Specht and team (Specht and Specht 1999a). 2.9.7. Chlorophyll a/b ratio (Carles Gracia on sabbatical from the University of Barcelona, Spain): This phenomenon was explored along the gradient from per-humid to arid climates Stewart et al. 1990). 2.9.8. Nitrate reductase enzyme activity (George Stewart on sabbatical from University College, London): Variation in this enzyme activity was explored along the gradient from per-humid to arid climates. If gaps occur in a plant community, solar radiation reaches the soil surface and increases the amount of nitrate in relation to ammonium ions formed during the decomposition of leaf litter (Stewart et al. 1990). Pioneer rainforest plants and exotic plants (weeds) — with thin leaves and high levels of nitrate reductase enzyme-activity — often invade these sunlit gaps (Stewart et al. 1988). The tall eucalypt trees (aka gigantism) that fringe rainforest pockets from north-eastern Australia to the Big Scrub of north-eastern New South Wales appear to possess thin leaves with high nitrate reductase enzyme-activity (Batianoff et al. 1997; Specht and Specht 1999a, p. 247). 2.9.9. Resurrection chloroplasts in the leaves of Arid Zone grasses (Cathy Mittelheuser, Dave Doley and Neil Trivett). With the onset of drought, the chloroplasts of the semi-arid-zone curly Mitchell grass (Astrebla lappacea) gradually shrink in size and their membrane structure changes substantially, but both structure (Mittelheuser 1977) and function (Doley and Trivett 1974) are largely restored within hours of re-wetting. Rod Rogers established that crustose lichens in the Arid Zone undergo similar changes during drying and wetting cycles (Rogers 1972). 2.9.10. Calcium carbonate toxicity (Bob Parsons in the University of Melbourne): — When sea levels fell during the last Ice Ages, the Australian continental shelf was exposed, leading to the weathering of biological remains, notably shells of molluscs and foraminifera fragments containing aragonite in southern Australia and massive coralline algal cell walls containing calcarenite, in north-eastern Australia. These calcareous dusts were blown inland and influenced vegetation development. Calcarenite inhibited the establishment of woody trees and shrubs on coastal dunes in northern Australia (Specht 1958); in southern Australia, the aragonite variously affected the different mallee species that occupied inland dunes (Parsons and Specht 1967). 2.8.11 Photosynthetic characteristics of Eucalyptus pauciflora (snow gum) along an altitudinal gradient (Ralph O. Slatyer of Australian National University): - The photosynthetic attributes of leaves of snow gum changed consistently along an altitudinal

20 gradient in the Australian Alps, indicating evolutionary adaptation to decreasing temperatures with increasing altitude (Slatyer 1977a, b, 1978; Slatyer and Ferrar 1977; Slatyer and Morrow 1977).

References Ashford, A. E., Ling-Lee, M. and Chilvers, G. A. (1975). Polyphosphate in eucalypt mycorrhizas: A cytochemical demonstration. New Phytologist 74: 447-453. Batianoff, G. N., Reeves, R. D. and Specht, R. L. (1997). The effect of serpentine on vegetation structure, species diversity and endemism in Central Queesland. Pp. 44-50. In: The Ecology of Ultramafic and Metalliferous Areas. (Eds T. Jaffré, R. D. Reeves and T. Becquer). Centre ORSTROM de Nouméa, New Caledonia. Brownell, P. F. (1965). Sodium as an essential micro-nutrient for a higher plant (Atriplex vesicaria). Plant Physiology 40: 460-468. Brownell, P. F. and Crossland, C. J. (1972). The requirement for sodium as a micro-nutrient by species having the C4 dicarboxylic photosynthetic pathway. Plant Physiology 49: 794-797. Burrell, Juliet P. (1981). Invasion of coastal heaths of Victoria by Leptospermum laevigatum (J. Gaertn.) F. Muell. Australian Journal of Botany, 29: 747-764. Coleman, R. G. and Specht, R. L. (1981). Mineral nutrition of heathlands. The possible role of polyphosphate in the phosphorus economy of heathland species. Pp. 197-207. In: Ecosystems of the World. Vol. 9B. Heathlands and Related Shrublands. Analytical Studies. (Ed. R. L. Specht). Elsevier, Amsterdam. Doley, D. and Trivett, N. B. A. (1974). Effects of low water potentials on transpiration and photosynthesis in Mitchell Grass (Astrebla lappacea). Australian Journal of Plant Physiology 1: 539-550. Farquhar, G. D., O’Leary, H. and Berry, J. A. (1982a). On the relationship between Carbon Isotope discrimination and the intercellular carbon dioxide concentration in leaves. Australian Journal of Plant Physiology 9: 121-137. Farquhar, G. D., B 13C value of halophytes. Evidence for diffusional isotope fractionation determined by the ratio of intercellular / atmospheric partial pressure of CO2 under different environmental conditions. Oecologia 52: 121-124. Jeffrey, D. W. (1964). The formation of polyphosphate in Banksia ornata, an Australian heath plant. Australian Journal of Biological Science 17: 845-854. Jeffrey, D. W. (1967). Phosphate nutrition of Australian heath plants. 1. The importance of proteoid roots in Banksia (Proteaceae). Australian Journal of Botany 15: 403-412 Jeffrey, D. W. (1968). Phosphate nutrition of Australian heath plants. II. The formation of polyphosphate by five heath species. Australian Journal of Botany 16: 603-613. Haines, Wendy M. (1967). Experimental Approaches to the Study of Seasonal Root Activity and Phosphorus Nutrition of Australian Heath Plants. M. Sc. Thesis, University of Melbourne. Hatch, M. D. and Slack, C. R. (1966). Comparative studies of the activity of carboxylases and other enzymes in relation to the new pathway of photosynthetic carbon dioxide fixation in tropical grasses. The Biochemical Journal 108: 660-665. Mittelheuser, Catherine, J. (1977). Rapid ultrastructural recovery of water stressed leaf tissue. Zeitschriff für Pflanzenphysiologie 82: 458-461. Ozanne, P. G. and Specht, R. L. (1981). Mineral nutrition of heathlands: Phosphorus toxicity. Pp. 209-213. In: Ecosystems of the World. Vol. 9B. Heathlands and Related Shrublands. Analytical Studies. (Ed. R. L. Specht). Elsevier, Amsterdam. Parsons, R. F. and Specht, R. L. (1967). Lime chlorosis and other factors affecting the distribution of Eucalyptus on coastal sands in southern Australia. Australian Journal of Botany 15: 95-105. Pate, J. S., Jeschke, W. D., Dawson, T. E., Raphael, C., Hartung, W. and Bowen, B. J. (1998). Growth and seasonal utilization of water and nutrients by Banksia prionotes. Australian Journal of Botany 46: 511-532. Rich, A. (1996). ‘What is Littoral about Rainforest?’ B. Appl. Sc. Hons. Thesis, Southern Cross University, Lismore, N.S.W. Rogers, R. W. (1972). Soil surface lichens in arid and sub-arid south-eastern Australia. II. Phytosociology and geographic zonation. III. The relationship between distribution and environment. Australian Journal of Botany 20: 215-227, 301-316. Slatyer, R. O. (1977a). Altitudinal variation in the photosynthetic characteristics of snow gum, Eucalyptus pauciflora Sieb. ex Spreng. IV. Temperature response of four populations grown at different temperatures. Australian Journal of Plant Physiology 4: 583-594. Slatyer, R. O. (1977b). Altitudinal variation in the photosynthetic characteristics of snow gum, Eucalyptus pauciflora Sieb. ex Spreng. VII. Comparison of field and phytotron responses to growth temperature. Australian Journal of Plant Physiology 4:901-916. Slatyer, R. O. (1978). Altitudinal variation in the photosynthetic characteristics of snow gum, Eucalyptus pauciflora Sieb. ex Spreng. XVI. Relationship between gradients of field temperature and photosynthetic optima in the Snowy Mountains area. Australian Journal of Botany 26:111-121. Slatyer, R. O. and Ferrar, P. J. (1977). Altitudinal variation in the photosynthetic characteristics of snow gum, Eucalyptus pauciflora Sieb. ex Spreng. I. Rate of acclimation to an altered growth environment. Australian Journal of Plant Physiology 4:595-609. Slatyer, R. O. and Morrow, P. A. (1977). Altitudinal variation in the photosynthetic characteristics of snow gum, Eucalyptus pauciflora Sieb. ex Spreng. I. Seasonal changes under field conditions in the Snowy Mountains area of south-eastern Australia. Australian Journal of Botany 25:1-20.

21 Specht, R. L. (1958). Climate, geology, soils and plant ecology of the northern potion of Arnhem Land. Pp. 415-478. In: Records of the American-Australian Scientific Expedition to Arnhem Land. Vol. 3.Botany and Plant Ecology. (Eds R. L. Specht & C. P. Mountford). Melbourne University Press, Melbourne. Specht, R. L. and Specht, Alison (1999, 2002). Australian Plant Communities. Dynamics of Structure, Growth and Biodiversity. Oxford University Press, Melbourne. Stewart, G. R., Hegarty, Elwyn E. & Specht, R. L. (1988). Inorganic nitrogen assimilation in plants of Australian rainforest communities. Physiologia Plantarum 74: 26-33. Stewart, G. R., Gracia, C. A., Hegarty, Elwyn E. and Specht, R. L. (1990). Nitrate reductase activity and chlorophyll content in sun leaves of subtropical Australian closed-forest (rainforest) and open-forest communities. Oecologia (Berlin) 82: 544-551.

3. Ecosystem Conservation

3.1. Classification of Australian plant communities Early in the 1950s, for the third edition of The Australian Environment, Prof. Joe Wood assembled ecologists (Fred Jessup, Ted Coaldrake, Ray Perry and Ray Specht) to prepare a two-way table to identify structural formations in the Australian vegetation. Raunkiaer’s Life Forms (approximate height) of the overstorey were used on the Y-axis. A crude estimate of the spatial distance between the canopies of the overstorey was used for the X-axis.

For the fourth edition, the soil scientist, Geoffrey Leeper invited Specht (1970) to write the chapter on ‘Australian Vegetation’. In preparation, a precise measurement of Foliage Projective Cover in the overstorey (> 4-6 m) was made using an upward-viewing crosswire sighting tube, at 1-m intervals along a line transect. The Foliage Projective Cover of the understorey (< 2 m) was measured using a downward- viewing crosswire sighting tube. The technique was developed from the pioneering work of Levy and Madden (1933), Tiver and Crocker (1951), Winkworth (1959) and Winkworth and Goodall (1962). This two-way classification of structural formations (closed-forest, open-forest, woodland, etc.) has been used to this day.

References Levy, E. B. & Madden, E. A. (1933). The point method of pasture analysis. New Zealand Journal of Agriculture 46: 267- 279. Specht, R. L. (1970). Vegetation. pp. 44-67 in The Australian Environment. 4th Edition. (Ed. G. W. Leeper) C.S.I.R.O. - Melbourne University Press, Melbourne. Tiver, N. S. & Crocker, R. L. (1951). The grasslands of south-eastern South Australia in relation to climate, soils and development history. Journal of the British Grassland Society 6: 29-80. Winkworth, R. E. (1955). The use of point quadrats for the analysis of heathland. Australian Journal of Botany 3: 68-81. Winkworth, R. E. & Goodall, D. W. (1962). Crosswire sighting tube for point quadrat analysis. Ecology 43: 342-343.

3.2 Conservation of Australian plant communities

In the late 1950s, the Australian Academy of Science fostered a ‘conservation survey’ of the major plant communities in each State and Territory. As ecologists had mapped most of the State of South Australia, Ray Specht and the 80-year-old Prof. J. B. Cleland assessed the conservation of plant communities throughout the State. As the flora of the State was also well documented, thanks to J. M. Black’s ‘Flora of South Australia’, they assessed the conservation state of the flora as well (Specht and Cleland 1961, 1964).

As Australian Convenor of the Conservation Section of the International Biological Program, Specht encouraged teams of ecologists to extend the South Australian survey to all States and Territories, led by:

Australian Capital Territory — Mrs Elizabeth Hayden

22 New South Wales — Dr A. B. Costin Northern Territory — Mr R. A. Perry Queensland — Prof. R. L. Specht & Mrs Ethel M. Roe South Australia — Mr C. W. Bonython Tasmania — Prof. W. D. Jackson Victoria — Mrs Judith Frankenberg & Prof. J. S. Turner Western Australia — Dr W. D. L. Ride Papua New Guinea — Dr D. N. McVean & Mr J. S. Womersley Australian Conservation Foundation — Dr F. N. Ratcliffe & Dr J. G. Mosley Australian Academy of Science — Dr M. F. Day

As this conservation survey was based on all the descriptive ecological surveys that had been made throughout Australia (Specht, Marion M. and Specht 1962), the Australian Heritage Commission encouraged the team to tabulate all the species information. This resulted in a tabulation of the conservation status of plant communities (Specht et al. 1974). Continuing analysis of data - on the mainframe computers in the University of Queensland and CSIRO — was undertaken to objectively define plant communities using TWINSPAN Floristic Groups and to determine their conservation status. This task took almost twenty years and resulted in a landmark quantitative evaluation of plant community conservation throughout Australia (Specht et al. 1995), summarised in Table 1. It has been followed by more extended analyses (Specht [Alison] and Specht 2000, 2013, 2016). The hazards of data collection and management of this and associated projects have been discussed recently by Specht [Alison] et al. (2018).

References Specht, Alison & Specht, R. L. (2000). Australia: Biodiversity of ecosystems. Pp. 307-324. In: Encyclopedia of Biodiversity. (Ed. S. Levin) Academic Press, San Diego, California. Specht, Alison & Specht, R. L. (2013). Australia: Biodiversity of ecosystems. Pp. 291-306. In: Encyclopedia of Biodiversity. Vol. 1. (Ed. S. Levin) Elsevier, Amsterdam. Specht, Alison & Specht, R. L. (2016). Australia: Biodiversity of ecosystems. In: Reference Module in Life Sciences. Elsevier, Amsterdam. Specht, Alison, Bolton, M. P., Kingsford, B., Belbin, L. & Specht, R. L. (2018). A story of data won, data lost and data re- found: The realities of ecological data. Biodiversity Data Journal 6: e28073 (doi: 10.3897/BDJ6.e28073). Specht, Marion M. and Specht, R. L. (1962). Bibliographia phystosociologica: Australia. Excerpta Botanica, Sectio B 4: 1- 58. Specht, R. L. and Cleland, J. B. (1961). Flora conservation in South Australia. 1. The preservation of plant formations and associations recorded in South Australia. Transactions of the Royal Society of South Australia 85: 177-196. Specht, R. L. and Cleland, J. B. (1964). Flora conservation in South Australia. 2. The preservation of species recorded in South Australia. Transactions of the Royal Society of South Australia 87: 63-92. Specht, R. L., Roe, E. M. & Boughton, V. H. (Eds) (1974). Conservation of Major Plant Communities in Australia and Papua New Guinea. Australian Journal of Botany Suppl. No. 7, 667 pp. Specht, R. L., Specht, Alison, Whelan, M. B. & Hegarty, E. E. (1995). Conservation Atlas of Plant Communities in Australia. Centre for Coastal Management, Southern Cross University, Lismore, N.S.W., 1073 pp.

23 TABLE 1. The conservation status (1995) of major TWINSPAN Floristic Groups (Hill 1973) that contain the majority of plants and resident animals in Australia (Analysed by M.P. Dale CSIRO Div. Computing Research). * Species that occurred only once in the TWINSPAN analysis were not included.

Formation/Complex Sampling Species* TWINSPAN Reasonable Collators Sites analysed Floristic conservation Groups extent

Rainforest 644 1,418 19 70% L.J.Webb, J.G. Tracey & R.L. Specht

Monsoon rainforest 1,219 559 17 31% J. Russell-Smith

Dry scrub (SE Qld) 232 475 17 26% P.D. Bostock, P.I. Forster & L.B. Hutley

Eucalypt forest 1,275 276 69 49% D.W. Drake & M.P. (trees only) Bolton

Sclerophyll (heath) 172 1,761 18 64% A. Specht & A.J.M. understorey (SW Hopkins W.A.)

Heathland/scrub 312 2,071 19 53% W.E. Drake, M.P. Bolton & R.L. Specht

Alpine heath & 110 556 16 67% J.B. Kirkpatrick grassland

Sclerophyll (heath) 372 2,581 21 34% R.L. Specht, W.E. Drake understorey & M.P. Bolton (eastern Aust.)

Grassland - savanna 200 1,313 27 13% M.P. Bolton, A. Specht understorey & R.L. Specht

Mallee (incl. SW 41 395 8 41% R.L. Specht & M.P. W.A.) Bolton

Desert Acacia (incl. 150 1,229 18 17% M.P. Bolton, A. Specht understorey of & R.L. Specht hummock grasses)

Hummock 26 303 10 30% R.J. Specht & E.E. grassland Hegarty

Chenopod 55 410 10 26% R.L. Specht & A. Specht shrubland

Forested wetland 77 835 18 24% R.L. Specht & A. Specht

Freshwater swamp 80 139 17 25% R.L. Specht & M.P. Bolton

Coastal dune 50 315 11 69% R.L. Specht & M.P. Bolton

Coastal wetland 15 74 12 47% R.L. Specht & A. Specht

24  Species-richness of vascular plants & resident vertebrates

Alpha biodiversity records the number of plant-species found in a plant community when the species-area curve asymptotes to a maximum — about 10 hectares in open-structured plant communities (Whittaker 1972); about 200 hectares in closed-structured (rainforest) plant communities (Specht A. 1988; Stocker 1988; Specht 2007).

Beta biodiversity (Whittaker 1972) measures the change in biodiversity of species from one plant community to another along an ecological gradient.

Gamma (community) biodiversity is the total biodiversity of all plant communities in a landscape, including both Alpha and Beta Biodiversities (Whittaker 1972) — especially high in the sclerophyll (heathy) and mallee communities in south-west Western Australia and in the rainforest stands in north-eastern Queensland.

3.3.1. Alpha Biodiversity (Species Richness) of vascular plants & resident vertebrates

Alpha diversity is correlated with the annual photosynthetic production of the Leaf Area Index (leaf area per unit ground area) and leaf characteristics, including mean individual leaf area, leaf specific weight (leaf weight per unit leaf area), and carbon isotope ratio. All of these attributes are related to the moisture index (Ea/Eo) and the Evaporative Coefficient of the location, and to the mean daily solar energy input, which decreases from the tropical north to the temperate south of the Australian continent (Specht A. and Specht 1993).

The edges of residual stands of rainforest (in well-watered ‘oases’ or in areas remaining after clearing) are subjected to a more desiccating environment and to more physical abrasion due to wind than is the interior of the forest. The combination of these effects reduces the annual photosynthesis and species- richness of plants, and also resident bird populations, in the stand.

The Alpha Biodiversity of closed-forests in Australian per-humid climates declines from about 150 species of overstorey trees per hectare in north-east Queensland until closed-forests disappear south of Tasmania, as solar radiation decreases (Stocker 1988; Hegarty 1991; Specht A. 1988; Rich 1996; Howard 1973). The Alpha Biodiversity of open-forests on medium-nutrient soils declines from about 10 species of overstorey eucalypts per hectare in south-east Queensland to one or two per hectare in Victoria and Tasmania (Specht and Perry 1948: Clifford 1954; Specht et al. 1991).

The number of species of lichens at or near ground level declined as more solar radiation reached the ground between the trees in open-structured plant communities in the gradient from the tropical north to the Tropic of Capricorn in Queensland (Rogers and Stevens 1981).

The high Alpha Biodiversity of birds on the black soil plain of the East Alligator River had been noted by Bert Deignan during the 1948 American-Australian Scientific Expedition to Arnhem Land (Deignan 1964), whereas the Alpha Biodiversity of plants was low on these seasonally waterlogged soils (Specht 1958), on which many invertebrates may thrive.

25 Prompted by the research conducted by CSIRO Division of Wildlife Research on mammals and plants in mature plant communities in northern Australia (Braithwaite et al. 1985) and by Martin Cody of the University of California Los Angeles on resident birds of the mulga of Central Australia (Cody 1994a,b), Specht and colleagues established that the Alpha diversities of plants and resident vertebrates (small mammals, birds, snakes, lizards and frogs) were correlated (Catling 1988; Specht and Specht 1999a, p. 332). It was hypothesised that the species richness of invertebrates that fell into pit-fall traps was also correlated.

The Alpha biodiversity of both plants and all resident vertebrates in mature plant communities — in humid to arid climates — in south-west Western Australia were lower than in south-eastern Australia (Specht and Specht 1999a, p. 332). It was hypothesised that this marked difference was probably due to the less fertile soils in south-west Western Australia. Specht and Tyler (2010) listed the Alpha biodiversity of vascular plants and Amphibia in Australian ecosystems found in localities with soils containing diverse levels of phosphate. When the annual shoot growth of each of these plant communities was estimated on a soil-nutrition curve, the diverse Alpha diversities all fell on the same line (Specht and Tyler 2019).

The species-richness of plots treated with and without phosphate in the experiment that was established on Dark Island heath was recorded in 1952, 1960, 1972 and 1989 (Heddle and Specht 1975; Specht and Specht 1999a, p. 342). It was noted that the understorey plants reached their peak of growth and died in a sequence termed post-fire succession. As the heath regenerated post-fire, the overstorey foliage-cover of Banksia ornata, B. marginata and Allocasuarina pusilla increased with time; the species-richness of the understorey declined (Specht and Specht 1999a, p.342) — faster on phosphate-treated than on control plots that experienced phosphorus-toxicity (Ozanne and Specht 1981).

It is apparent that a bush-fire is necessary to restore the biodiversity of this plant community.

References Batianoff, G. N. and Specht, R. L. (1992). Queensland (Australia) serpentine vegetation. Pp. 109-128. In: The Vegetation of Ultramafic (Serpentine) Soils. (Eds A. J. M. Baker, J. Proctor and R. D. Reeves). Intercept Publications, Andover, England. Batianoff, G. N., Specht, R. L. and Reeves, R. D. (1991). The serpentinite flora of the humid subtropics of eastern Australia. Proceedings of the Royal Society of Queensland 101: 137-158. Braithwaite, R. W., Winter, J. W., Taylor, J. A. & Parker, B. S. (1985). Patterns of diversity and structure of mammalian assemblages in the Australian tropics. Australian Mammalogist 8: 171-186. Catling, P. C. (co-ordinator) (1988). Vertebrates. Pp. 171-194. In: Mediterranean-type Ecosystems. A Data Source Book. (Ed. R. L. Specht) Kluwer Academic Publishers, Dordrecht. Clifford, H. T. (1954). On the distribution of the species of Eucalyptus in the region of the Dandenong Range, Victoria. Proceedings of the Royal Society of Victoria 65: 449-455. Cody, M. L. (1994a). Bird diversity within and among heathlands. Pp. 47-61. In: Plant-Animal Interactions in Mediterranean-type Ecosystems. (Eds Margarita Arianoutsou & R. H. Groves) Kluwer Academic Publishers, Dordrecht. Cody, M. L. (1994b). Mulga bird communities. I. Species composition and predictability across Australia. Australian Journal of Ecology 19: 206-219. Deignan, H. G. (1964). Birds of the Arnhem Land Expedition. Pp. 345-425. In: Records of the American-Australian Scientific Expedition to Arnhem Land. Vol. 4. Zoology. (Ed. R. L. Specht). Melbourne University Press, Melbourne. Heddle, Elizabeth M. and Specht, R. L. (1975). Dark Island heath (Ninety-Mile Plain, South Australia). 8. The effect of fertilizers on composition and growth, 1950-1972). Australian Journal of Botany 23: 151-164. Hegarty, Elwyn E. (1991). Patterns of productivity by liane and tree components of an Australian rainforest. Journal of Tropical Ecology 7: 201-214. Howard, Truda M. (1973). Studies in the ecology of Nothofagus cunninghamii Oerst. II. Phenology. Australian Journal of Botany 21: 79-92.

26 Ozanne, P. G. & Specht, R. L. (1981). Mineral nutrition of heathlands: Phosphorus toxicity. Pp. 209-213. In: Ecosystems of the World. Vol. 9B. Heathlands and Related Shrublands. Analytical Studies. (Ed. R. L. Specht) Elsevier, Amsterdam. Rich, A. (1996). ‘What is Littoral about Rainforest?’ B. Appl. Sc. Hons. Thesis, Southern Cross University, Lismore. Rogers, R. W. & Stevens, G. Nell (1981). Lichens. Pp. 591-603. In: Ecological Biogeography of Australia. (Ed. A. Keast) Junk, The Hague. Specht, Alison (1988). Big Scrub Conservation Strategy. Vol. 2. Resource Material. (Ed. Planners North Pty Ltd) Pp. 2.1-2,72. New South Wales National Parks and Wildlife Service, Sydney. Specht, Alison & Specht, R. L. (1993). Species richness and canopy productivity of Australian plant communities. Biodiversity and Conservation 2: 152 - 167. Specht, Alison & Specht, R. L. (1994). Biodiversity of overstorey trees in relation to canopy productivity and stand density in the climatic gradient from warm temperate to tropical Australia. Biodiversity Letters 2: 39-45. Specht, R. L. (1958). Climate, geology, soils and plant ecology in the northern portion of Arnhem Land. Pp. 333- 414. In Pp. 345-425. In: Records of the American-Australian Scientific Expedition to Arnhem Land. Vol. 3. Botany and Plant Ecology. (Eds R. L. Specht and C. P. Mountford). Melbourne University Press, Melbourne. Specht, R. L. (2007). Species richness of rainforest stands on non-serpentinite and serpentinite substrates in the Rockhampton – Marlborough area of Central Queensland. Proceedings of the Royal Society of Queensland 113: 17-35. Specht, R. L. and Perry, R. A. (1948). Plant ecology of part of the Mount Lofty Ranges. Transactions of the Royal Society of South Australia 72: 91-132. Specht, R.L. & Specht, Alison (1989). Species richness of overstorey strata in Australian plant communities. The influence of overstorey growth rates. Australian Journal of Botany 37: 321-336. Specht, R. L. & Specht, Alison (1999, 2002). Australian Plant Communities. Dynamics of Structure, Growth and Biodiversity. Oxford University Press, Melbourne. Specht, R. L. & Specht, Alison (2007). Pre-settlement tree density in the eucalypt open-forest on the Brisbane Tuff. Proceedings of the Royal Society of Queensland 113: 9-16. Specht, R. L. & Specht, Alison (2013). Australia: Biodiversity of ecosystems. Pp.291-306. In: Encyclopedia of Biodiversity. (Ed. S. Levin). Elsevier, Amsterdam. Specht, R. L. & Tyler, M. J. (2010). The species richness of vascular plants and Amphibia in major plant communities in temperate and tropical Australia: relationship with annual biomass production. International Journal of Ecology 2010, Article ID 635852, 17 pages. Specht, R. L., Clifford, H. T., Arianoutsou, Margarita, Bird, L. H., Bolton, M. P., Forster, P. I., Grundy, Rhonda I., Hegarty, Elwyn E. and Specht, Alison (1991). Structure, floristics and species richness of plant communities in south-east Queensland. Proceedings of the Royal Society of Queensland 101: 27-78. Stocker, G. C. (1988). Tree species diversity in rainforests — establishment and maintenance. Proceedings of the Ecological Society of Australia 15: 39-47. Whittaker, R. H. (1972). Evaluation and measurement of species diversity. Taxon 21: 213-251.

3.3.2. Beta Biodiversity (Species Richness along ecological gradients)

Beta biodiversity (Whittaker 1972) measures the change in biodiversity of species from one plant community to another along an ecological gradient.

Beta Biodiversity was studied in an eucalypt open-forest as soil depth increased in the serpentinite outcrop in Central Queensland (Batianoff and Specht 1992; Batianoff et al. 1991). The density of the overstorey trees (FPCo) increases as soil depth (and water retention) increases, thus decreasing the amount of solar radiation reaching the understorey strata. As the Foliage Projective Cover of the overstorey (FPCo) increases, the Foliage Projective Cover of the understorey (FPCu) decrease FPCu) remains constant. The component species along this edaphic gradient provides a measure termed Beta Biodiversity of this edaphic gradient.

The sequence of changes (post-fire) in the overstorey and understorey of the Dark Island heathland in the Upper South East of South Australia shows Beta Biodiversity — with FPCo increasing in the overstorey over 26 years (post-fire), causing a continuous decline in FPCu and species richness in the understory, while the sum Σ (FPCo + FPCu) remains constant (Specht 1963; Heddle and Specht 1975; Specht and Specht 1999a, p. 342). The component species along this post-fire gradient provides a measure termed Beta Biodiversity.

27

References Batianoff, G. N. and Specht, R. L. (1992). Queensland (Australia) serpentine vegetation. Pp. 109-128. In: The Vegetation of Ultramafic (Serpentine) Soils. (Eds A. J. M. Baker, J. Proctor and R. D. Reeves). Intercept Publications, Andover, England. Batianoff, G. N., Specht, R. L. and Reeves, R. D. (1991). The serpentinite flora of the humid subtropics of eastern Australia. Proceedings of the Royal Society of Queensland 101: 137-158. Heddle, Elizabeth M. and Specht, R. L. (1975). Dark Island heath (Ninety-Mile Plain, South Australia). 8. The effect of fertilizers on composition and growth, 1950-1972). Australian Journal of Botany 23: 151-164. Specht, R. L. (1963). Dark Island heath (Ninety-Mile Plain, South Australia). 8. The effect of fertilizers on composition and growth, 1950-1960.. Australian Journal of Botany 11: 67-94. Specht, R. L. & Specht, Alison (1999, 2002). Australian Plant Communities. Dynamics of Structure, Growth and Biodiversity. Oxford University Press, Melbourne. Whittaker, R. H. (1972). Evaluation and measurement of species diversity. Taxon 21: 213-251.

3.3.3. Gamma Biodiversity (Species Richness per region)

The high Gamma Biodiversity of south-west Western Australia probably resulted from the severe drought conditions that occurred in south-western Australia when Global Warming occurred about ten million years ago (Martin 2006). The vegetation was then confined to well-watered ‘oases’ where evolution could occur. When a more favourable climate returned, the plants in these ‘oases’ recolonised the area forming plant communities with Alpha Biodiversity typical of the locality (George et al. 1979; Lamont et al. 1984; Specht and Specht 1989). The Gamma Biodiversity of the region was markedly increased.

A similar ‘disaster’ occurred in north-eastern Australia during the two major Ice Ages in the last million years (Kershaw 1981, 1992: Hopkins et al. 1993, 1996). Sea level fell, thus exposing the Great Barrier Reef. Severe droughts confined the rainforest to well-watered ‘oases’, while eucalypt forests invaded. When a favourable climate was restored, the new species in these ‘oases’ expanded to form the present rainforests of the region, thus creating a high Gamma Biodiversity in the region.

References Baker, G. (1959). 1. Opaline phytoliths in some Victorian soils and ‘red rain’ residues. 2. A contrast in the opaline assemblages in two Victorian soils. Australian Journal of Botany 7: 64-87, 88-96. Baker, G. (1960). Phytoliths in some Australian dusts. Proceedings of the Royal Society of Victoria 72: 21-40. Hopkins, M. S., Ash, J., Graham, A. W., Head, J. and Hewett, R. K. (1993). Charcoal evidence of the spatial extent of the Eucalyptus woodland expansions and rainforest contractions in North Queensland during the Late Pleistocenr. Journal of Biogeography 20: 357-372. Hopkins, M. S., Head, J., Ash, J. E., Hewett, R. K. and Graham, A. W. (1996). Evidence of a Holocene and continuing expansion of lowland rainforest in humid, tropical North Queensland. Journal of Biogeography 23: 737-745. Kershaw, A. P. (1981). Quaternary vegetation and environments. Pp. 81-101. In: Ecological Biogeography of Australia. (Ed. A. Keast). Junk, The Hague. Kershaw, A. P. (1992). The development of rainforest-savanna boundaries in tropical Australia. Pp. 255-271. In: Nature and Dynamics of Forest-Savanna Boundaries. (Eds P. A. Furley, J. Proctor & J. A. Ratter). Chapman & Hall, London. George, A. S., Hopkins, A. J. M. and Marchant, N. G. (1979). The heathlands of Western Australia. Pp. 211-230. In: Ecosystems of the World. Vol. 9A. Heathlands and Related Shrublands. Descriptive Studies. (Ed. R. L. Specht). Elsevier, Amsterdam. Lamont, B. B., Hopkins, A. J. N. and Hnatiuk, R. J. (1984). The flora — composition, diversity and origins. Pp. 27-50. In: Kwongan: Plant Life of the Sandplain. (Eds J. S. Pate and J. S. Beard). University of Western Australia Press, Nedlands. Martin, Helene A. (2006). Cainozoic climatic change and the development of the arid vegetation in Australia. Journal of Arid Enviroments. 66: 533-563. Specht, R. L. and Specht, Alison (1989). Species richness of sclerophyll (heathy) plant communities in Australia. The influence of overstorey cover. Australian Journal of Botany 37: 321-336.

28 3.4 Seed dispersal

Seed dispersal mechanisms of the genera in 16 plant formations in Australia were assessed by Trevor Clifford and Ray Specht using the following characteristics (Clifford & Specht 1999, pp. 20-23): —

Anemochory (dispersal by wind currents)

Hydrochory (flotation on or movement in bodies of water)

Censer (slow release of seed from the fruit by shaking in the wind)

Ballistochory (explosive release of seed from the fruit)

Epizoochory (on the fur, feathers, etc. of animals)

Endozoochory (passing through the digestive tract of animals)

Cache zoochory including myrmecochory (harvesting by ants)

Some 50% of seeds in a subtropical rainforest are fleshy and eaten by birds and bats that distribute the seed nearby. An abandoned farm adjoining the Mt Glorious rainforest, near Brisbane, had been re- established with all nearby rainforest species after 50 years (Hegarty 1988). Similarly, an abandoned paddock at O’Reilly’s, adjoining Lamington National Park, had been almost regained its original species richness after 50 years (Hopkins 1975).

The seeds of eucalypts (censers) are shaken from their capsules and drift in the wind barely more than twice the height of the tree. Understorey plants lacking specialised seed distribution mechanisms move even smaller distances. However, over thousands of years, complete communities of heathy woodlands have occupied the sandy off-shore islands in south-eastern Queensland — South and North Stradbroke Islands, Moreton Island, Bribie Island, Cooloola, and Fraser Island — probably spreading from small stands of heathy vegetation around rocky outcrops nearby.

References Clifford, H. T. & Specht, R. L. (1999). Seed dispersal. Pp. 20-23. In: Australian Plant Communities. Dynamics of Structure, Growth and Biodiversity. (Specht, R. L. & Specht, Alison). Oxford University Press, Melbourne. Hegarty, Elwyn E. (1988). Canopy Dynamics of Lianes and Trees in Subtropical Rainforest. Ph. D. Thesis, University of Queensland.

Hopkins, M. S. (1975). Species Patterns and Diversity in the Subtropical Rain Forest. Ph. D. Thesis, University of Queensland.

3.5 Species evolution in seepage of uraniferous radiation

Early research at the Atomic Research Station at Chalk River, Canada, showed that background ionising radiation produced increasing genetic changes in the eyes of trout whose sperms have been exposed to radiation from zero up to 400 rads (McGregor & Newcombe 1972).

During the American-Australian Scientific Expedition to Arnhem Land in 1948, samples of uni-cellular algae associated with pond-weed were collected from the freshwater Unawalerke Lagoon near the Arnhem Land escarpment at Oenpelli (Gunbalanya), Northern Territory. This collection contained 164 desmid taxa (48 undescribed), far richer in taxa than other samples (10-25 taxa) collected elsewhere in the Northern Territory, South Australia or overseas (Prescott & Scott 1952; Scott & Prescott 1958; Croasdale & Scott 1976).

29 In more recent work, Phreatoicidean isopods show ‘micro-endemism’ in Arnhem Land, with one or two unique taxa each with an apparent range of less than one kilometre (Wilson et al. 2009). South of Gunbalanya, Wilson found three new species and in the North Jabiluka outlier, there were as many as 11 species.

The seasonally waterlogged, infertile sandy soils in the humid to sub-humid climate of the Northern Territory support a diverse assemblage of short-lived plant species (Cowie 2005):

Utricularia with 36 taxa (Taylor 1989); Eriocaulon with 21 taxa (Leach 2000); Lindernia with over 25 taxa (W. R. Barker, unpublished)

Byblis, Calandrinia, Centolepis, Oldenlandia, Stylidium, Typhonium also show speciation. In contrast to these short-lived plants, perennial plants near uranium deposits in the Top End do not show any speciation (Bill Bateman, pers. comm. Rum Jungle: Ray Specht 1958b, 2012).

References Cowie, I. D. (2005). Goodenia elaiosoma I. D. Cowie (Goodeniaceae), a new species from the Top End of the Northern Territory and a key to the northern species. Austrobaileya 7: 205-213. Croasdale, H. & Scott, A. M. (1976). New or otherwise interesting desmids from Northern Australia. Nova Hedwigia 27: 591-595. McGregor, J. F. & Newcombe, H. B. (1972). Dose-response relationships for yields of major eye malfunctions following low doses of radiation in trout sperm. Radiation Research 49: 155-169. Prescott, G. W. & Scott, A. M. 1952). Some South Australian desmids. Transactions of the Royal Society of South Australia 75: 55-69. Scott, A. M. & Prescott, G. W. (1958). Some freshwater algae from Arnhem Land in the Northern Territory of Australia. Pp. 9-136. In: Records of the American-Australian Scientific Expedition to Arnhem Land. Vol. 3.Botany and Plant Ecology. (Eds R. L. Specht & C. P. Mountford). Melbourne University Press, Melbourne. Specht, R. L. (1958). Climate, geology, soils and plant ecology of the northern potion of Arnhem Land. Pp. 415-478. In: Records of the American-Australian Scientific Expedition to Arnhem Land. Vol. 3.Botany and Plant Ecology. (Eds R. L. Specht & C. P. Mountford). Melbourne University Press, Melbourne. Specht, R. L. (2012). American-Australian Scientific Expedition to Arnhem Land (1948): Its long-range impact. The Open Ecology Journal 5: 53-83. Wilson, G. D. F., Humphrey, C. L., Colgan, D. J., Gray, K-A. & Johnson, R. N. (2009). Monsoon-influenced speciation patterns of a species flock of Eophreatoicus Nicholls (Isopoda: Crustacea). Molecular Phylogenetics and Evolution 51: 349-364.

3.6 Palaeo-botanical studies in Australia

During 1958, the President of the Royal Society of South Australia, Dr C. G. Stephens of CSIRO Soils Division, formed a workshop to promote scientific research on the present landscapes of Australia.

CSIRO Soils Division promoted the study of the geomorphology of sand-dunes that had been formed in many coastal locations around the continent with fluctuations in sea-levels during the Quaternary.

As a member of this committee, Ray Specht promoted the use of palynology in the study of the history of Australian vegetation, from the Cretaceous to the Recent (see Hill 20017).

Four major areas of research were developed: climate and soils; Late Cretaceous palynology; Mid-Tertiary palynology; Quaternary palynology and vegetation.

3.6.1 Climate and Soils

In the Late Cretaceous, a tropical to subtropical climate existed throughout the Gondwanan super- continent. Lateritic podsolic soils had been developed on the peneplain surfaces of all the continents of

30 Gondwanaland during the Late Cretaceous (Prescott and Pendleton 1952). The kaolonitic clay that had been developed in the subsoil fixed phosphate and molybdenate ions within its clay lattice — making these elements unavailable to pasture plants (Williams and Andrew 1970); somehow heathland plants and associated eucalypts had managed to survive on these extremely infertile soils (Specht and Perry 1948; Specht et al. 1958; Coleman and Specht 1981).

Over the millennia, the peneplain landscape of Australia was changed by the uplift of mountain ranges; the kaolonite clay of the Cretaceous lateritic podsols was included as an erosion product in many Present-Day soils (Stace et al. 1968), where it still plays a threat to agriculture (Specht 2001, 2002).

Dr Chris Stephens of CSIRO Soils Division pursued his analyses of pedogenesis following the dissection of lateritic regions in southern Australia. (Stephens 1946) with a study of laterites and silcretes in Australia (Stephens 1971).

The continental plates that formed Gondwanaland began to drift apart during the Tertiary. The Australian Plate (including Australia and New Guinea) separated from Antarctica and drifted northward at an average of 66 mm per year — according to the dating of lava-flows that were produced when the continent drifted over a ‘hot-spot’ in the Earth’s crust (Wellman and McDougall 1974). The formation of the Southern Ocean enabled the Atlantic currents to flow into the Pacific and across to South America. The climate of Australia became gradually cooler with a seasonal Mediterranean-type climate in the south. See Figure 1.

Figure 1: Distribution of lateritic soils on the continental land masses of Gondwanaland (after Prescott & Pendleton 1952 in Specht & Specht 1999).

31 Based on the isotopic signature of δ18O in kaolinitic soil profiles, Bird et al. (1990) suggested that a rise in atmospheric carbon dioxide was associated with these weathering profiles during both the Late Cretaceous and the Mid Tertiary.

Global Warming occurred, again, about ten million years ago and lead to the production of some lateritic podsols in Central Queensland. However, the lateritic podsols of northern Australia, even in south-west Western Australia, were further leached to form lateritic earths — sometimes containing bauxite (Specht 1958b; Specht et al. 1977). The silica in the leaf-litter under the desert Spinifex (Triodia spp.) was leached to form silcretes (Baker 1959, 1960; Stephens 1971) — some of which flowed down-stream to form the Sturt Stony Desert. Sea-level rose and swamped the Nullarbor Plain and the Murray-Darling Basin — creating deep limestone deposits in both these basins.

The fluctuations in sea-level during the Ice Ages of the Quaternary (Shackleton and Kennett 1975; Bird et al. 1990; Feary et al. 1991) enabled great amounts of sand to be blown onto the coast in south-west Western Australia, South Australia and south-east Queensland. In south-eastern South Australia, a slight rise in the earth’s crust enabled the survival of about ten coastal dunes (Crocker 1944, 1946; Schwebel 1984) from which the sand was blown easterly to form the Ninety-Mile Plain in South Australia, the Little and Big Deserts in Victoria. In Queensland, sand built up behind rocky outcrops to form the dune systems on Fraser Island, Cooloola, Bribie Island, Moreton Island and North and South Stradbroke Island. At Cooloola, six layers of overlapping dunes have been discovered. Along this sequence, the vegetation gradually develops into a eucalypt open-forest by Dune-System 4, dated at 90,000 years. Leaching in older Dune Systems (over 730,000 years old) has gradually become unfavourable for the growth of trees, with only a heathland surviving (Thompson 1981, 1992; Tejan-Kella et al. 1990; Tibby et al. 2017).

The high Gamma Biodiversity of south-west Western Australia probably resulted from the severe drought conditions that occurred in south-western Australia when Global Warming occurred about ten million years ago. The vegetation was then confined to well-watered ‘oases’ where evolution could occur. When a more favourable climate returned, the plants in these ‘oases’ recolonised the area forming plant communities with Alpha Biodiversity typical of the locality. The Gamma Biodiversity of the region was markedly increased.

A similar ‘disaster’ occurred in north-eastern Australia during the two major Ice Ages in the last million years. Sea level fell, thus exposing the Great Barrier Reef. Severe droughts confined the rainforest to well- watered ‘oases’, while eucalypt forests invaded. When a favourable climate was restored, the new species in these ‘oases’ expanded to form the present rainforests of the region, thus creating a high Gamma Biodiversity in the region.

References Baker, G. (1959). 1. Opaline phytoliths in some Victorian soils and ‘red rain’ residues. 2. A contrast in the opaline assemblages in two Victorian soils. Australian Journal of Botany 7: 64-87, 88-96. Baker, G. (1960). Phytoliths in some Australian dusts. Proceedings of the Royal Society of Victoria 72: 21-40. Bird, M. L., Fyfe, B., Chivas, A. & Longstaff, F. (1990). Deep weathering at extra-tropical latitudes: a response to increased atmospheric CO2. Pp. 383-389. In: Proceedings of the International Conference, Soils and the Greenhouse Effect. (Ed. A. F. Bouwman) Wiley, Chichester.

32 Coleman, R. G. and Specht, R. L. (1981). Mineral nutrition of heathlands: The possible role of polyphosphate in the phosphorus economy of heathland species. Pp. 197-207. In: Ecosystems of the World. Vol. 9B. Heathlands and Related Shrublands. Analytical Studies. Elsevier, Amsterdam. Crocker, R. L. (1944). Soil and vegetation relationships in the Lower South-East of South Australia. A study in ecology. Transactions of the Royal Society of South Australia 68: 144-172. Crocker, R. L. (1946). Post-Miocene Climatic and Geologic History and its Significance in Relation to the Major Soil Types of South Australia. Bulletin of the Council of Scientific and Industrial Research No. 193. Feary, D. A., Davies, P. J., Pigram, C. J. & Symonds, P. A. (1991). Climatic evolution and control of carbonate deposition in north-east Australia. Palaeogeography, Palaeoclimatology, Palaeoecology (Global and Planetary Change Section) 89: 341-361. Hill, R. S. (Ed.) (2017). History of the Australian Vegetation: Cretaceous to Recent. University of Adelaide Press, Adelaide. Kershaw, A. P. (1992). The development of rainforest-savanna boundaries in tropical Australia. Pp. 255-271. In: Nature and Dynamics of Forest-Savanna Boundaries. (Eds P. A. Furley, J. Proctor & J. A. Ratter). Chapman & Hall, London. Martin, Helene A. (2006). Cainozoic climatic change and the development of the arid vegetation in Australia. Journal of Arid Enviroments. 66: 533-563. Prescott, J. A. & Pendleton, R. L. (1952). Laterite and Lateritic Soils. Commonwealth Bureau of Soil Science and Technology, Communication No. 47, 51 pp. Schwebel, D. A. (1984). Quaternary stratigraphy and sea-level variation in the South East of South Australia. Pp. 291- 311. In: Coastal Geomorphology in Australia. (Ed. B. G. Thom). Academic Press Australia, Sydney. Shackleton, N. J. & Kennett, J. P. (1975). Palaeo-temperature history of the Cenozoic and the initiation of Antarctic glaciation: Oxygen and carbon isotope analyses in DSDP sites 277, 279, 281. Pp. 743-755. In: Initial Reports of the Deep Sea Drilling Project, 29. United States Government Printing Office, Washington. Specht, R. L. (1958). Climate, geology, soils and plant ecology of the northern portion of Arnhem Land. Pp. 333-414. In: Records of the American-Australian Scientific Expedition to Arnhem Land. Vol. 3. Botany and Plant Ecology. (Eds R. L. Specht and C. P. Mountford). Melbourne University Press, Melbourne. Specht, R. L. (2001). Phosphorus toxicity and pollution: A threat to our Gondwanan heritage. Ecological Management and Restoration 2: 228-230. Specht, R. L. (2002). Phosphate pollution and soil nitrate: Threats to biodiversity in Australia. Pp. 53-70, In: Landscape Health of Queensland. (Eds A. J. Franks, Julia Playford and A. Shapcott) Royal Society of Queensland, Brisbane. Specht, R. L. and Perry, R. A. (1948). . Plant ecology of part of the Mount Lofty Ranges. Transactions of the Royal Society of South Australia 72: 91-132. Specht, R. L., Rayson, Patricia and Jackman, Margaret E. (1958). Dark Island heath (Ninety-Mile Plain, South Australia). 6. Pyric succession: Changes in composition, coverage, dry weight and mineral nutrient status. Australian Journal of Botany 6: 50-88. Specht, R. L., Salt, R. B. and Reynolds, Sally (1977). Vegetation in the vicinity of Weipa, North Queensland. Proceedings of the Royal Society of Queensland 88: 17-38. Stace, H. C. T., Hubble, G. D., Brewere, R., Northcote, K. H., Sleeman, J. R., Mulcahy, M. J. and Hallsworth, E. G. (1968). A Handbook of Australian Soils. Rellim Technical Publications, Glenside, South Australia. Stephens, C. G. (1946). Pedogenesis following the dissection of lateritic regions in southern Australia. Bulletin of the Council of Scientific Research No. 206. Stephens, C. G. (1971). Laterite and silcrete in Australia. Geoderma 5: 5-52. Tejan-Kella, M. S., Chittleborough, D. J., Fitzpatrick, R. W., Thompson, C. H., Prescott, J. R. & Hutton, J. T. (1990). Thermo- luminescence dating of coastal sand dunes at Cooloola and North Stradbroke Island, Queensland, Australia. Australian Journal of Soil Research 28:465-481. Thompson, C. H. (1981). Podzol chronosequences in coastal dunes of eastern Australia. Nature 291: 59-61. Thompson, C. H. (1992). Genesis of podsols on coastal dunes in southern Queensland. I. Field relationships and profile morphology. Australian Journal of Soil Research 30: 593-613. Tibby, J., Barr, C., Marshall, J. C., McGregor, G. B., Moss, P. T., Arnold, L. J., Page, T. J., Questiaux, D, Olley, J., Kemp, J., Spooner, N., Petherick, L., Penny, D., Mooney, S. & Moss, E. (2017). Persistence of wetlands on North Stradbroke Island (south-east Queensland, Australia) during the last glacial cycle: implications for Quaternary science and biogeography. Journal of Quaternary Science 32: 770-781. Wellman, P. & McDougall, I. (1974). Cainozoic igneous activity in eastern Australia. Tectonophysics 23: 49-65. Willams, C. H. and Andrew, C. S. (1970). Mineral nutrition of pastures. Pp. 321-328. In: Australian Grasslands. (Ed. R. M. Moore). Australian National University Press, Canberra.

3.6.2 Late Cretaceous palynology

Mary Dettmann, who had studied with Dr Isabel Cookson, the pioneer of palynology of the lignite deposits in Victoria in the 1940s (Cookson 1954), furthered the study of the Late Cretaceous – Early Tertiary palynology of southern Australia and South America, a period when these continents were still attached to Antarctica (Dettmann 1981, 1989, 1994, 1998; Dettmann and Jarzen 1991, 1996 1998; Dettmann et al. 1995; Specht and Dettmann 1995; Specht et al. 2015). This research was summarised in ‘The History of Australian Vegetation: Cretaceous to Recent ‘ (Hill 2007).

33 The climate of southern Australia, at latitude 60oS was subtropical, with solar rays reaching this latitude throughout the year. A tall open-forest, similar to that in north-eastern New South Wales, was calculated to have existed in this environment (Specht et al. 1992).

References Cookson, Isobel C. (1954). The Cainozoic occurrence of Acacia in Australia. Australian Journal of Botany 2: 52-59. Dettmann, Mary E. ((1981). The Cretaceous flora. Pp. 355-375. In: Ecological Biogeography of Australia. (Ed. A. Keast) Junk, The Hague. Dettmann, Mary E. (1989). Antarctica: Cretaceous cradle of austral temperate rainforests? Pp. 89-105. In: Orogins and Evolution of the Antarctic Biota. (Ed. J. A. Crane) Geological Society of London Special Publication No. 47, London. Dettmann, Mary E. (1994). Cretaceous vegetation: The micro-fossil record. Pp. 143-170. In: History of the Australian Vegetation. (Ed. R. S. Hill) Cambridge University Press, Cambridge. Dettmann, Mary E. (1998). Patterns of radiation and diversification of some Gondwanan angiosperm families. Journal of African Earth Sciences 27: 60-61. Dettmann, Mary E. (2017). Cretaceous vegetation: The microfossil record. Pp. 143-170. In: History of Australian Vegetation: Cretaceous to Recent. (Ed. R. S. Hill). University of Adelaide Press, Adelaide. Dettman, Mary E. & Jarzen, D. M. (1991). Pollen evidence for Late Cretaceous differentation of Proteaceae in southern polar forests. Canadian Journal of Botany 69: 901-906. Dettman, Mary E. & Jarzen, D. M. (1996). Pollen of proteaceous-type from latest Cretaceous sediments, south-eastern Australia. Alcheringa 20: 103-160. Dettman, Mary E. & Jarzen, D. M. (1998). The early history of the Proteaceae in Australia: the pollen record. Australian Systematic Botany 11: 401-438. Dettman, Mary E., Jarzen, D. M. & Jarzen, S. A. (1995). Feeding habits of the mahogany glider; palynological evidence. Palynology 19: 137-142. Hill, R. S. (Ed.) (2007). History of the Australian Vegetation: Cretaceous to Recent. University of Adelaide Press, Adelaide. Specht, R.L. and Dettmann, M.E. (1995). Palaeo-ecology of Australia and current physiological functioning of plant communities. pp. 201-214. In: Time Scales of Biological Responses to Water Constraints: The Case of Mediterranean Biota. (Eds J. Roy, J. Aronson and F. di Castri) SPB Academic Publishing, Amsterdam. Specht, R. L., Dettmann, Mary E. & Jarzen, D. M. (1992). Community associations and structure in the Late Cretaceous vegetation of south-east Australasia and Antarctica. Palaeogeography, Palaeoclimatology, Palaeoecology 94: 283- 309. Specht, R. L., Montenegro, G. and Dettmann, M. E. (2015). Structure and alpha biodiversity of major plant communities in Chile, a distant biogeographical relation with Australia. Macrothink Institute Journal of Environment & Ecology 6: 21-47.

3.6.3 Mid-Tertiary palynology

Helene Martin, a post-graduate student who had studied the water-balance of eucalypt open-forest to woodland in the Mount Lofty Ranges, South Australia (Martin and Specht 1962, 2005), decided to train in palynology for her Ph. D. in Vancouver, Canada.

On completion of her Ph. D., she was appointed to study the bore-logs of the Murray-Darling Basin in New South Wales — based in the University of New South Wales. Later, her studies extended across the Arid Zone of Australia to the Indian Ocean.

Her observations indicated that Global Warming had occurred during the Mid-Tertiary (about 10 million years ago) and induced a long period of drought from the Pilbara across Australia into south-west New South Wales and north-west Victoria. This dry climate affected the vegetation of south-west Western Australia where inland heathy vegetation survived only in well-watered ‘oases’.

The high Gamma Biodiversity of south-west Western Australia probably resulted from the severe drought conditions that occurred in south-western Australia when Global Warming occurred about ten million years ago. The vegetation was then confined to well-watered ‘oases’ where evolution could occur. When a more favourable climate returned, the plants in these ‘oases’ recolonised the area forming plant communities with Alpha Biodiversity typical of the locality (George et al. 1979). The Gamma Biodiversity of the region had been markedly increased.

34 References George, A. S., Hopkins, A. J. M. and Marchant, N. G. (1979). The heathlands of Western Australia. Pp. 211-230. In: Ecosystems of the World. Vol. 9A. Heathlands and Related Shrublands. Descriptive Studies. (Ed. R. L. Specht). Elsevier, Amsterdam. Martin, Helene A. (1977). The Tertiary stratigraphy of the Murray Basin in New South Wales. 1. The Hay-Balranald- Wakool Districts. Journal and Proceedings of the Royal Society of New South Wales 110: 41-47. Martin, Helene A. (1978). Evolution of the Australian flora and vegetation through the Tertiary: Evidence from pollen. Alcheringa 2: 181-202. Martin, Helene A. (1981). The Teriary flora. Pp. 391-406. In: Ecological Biogeography of Australia. (Ed. A. Keast) Junk, The Hague. Martin, Helene A. (1996). Wildfires in past ages. Proceedings of the Linnean Society of New South Wales 116: 1-18. Martin, Helene A. (1987). Cainozoic history of the vegetation and climate of the Lachlan River region, New South Wales. Proceedings of the Linnean Society of New South Wales 109: 214-257. Martin, Helene A. (1990a). Teriary climate and phytogeography of south-eastern Australia. Review of Palaeobotany and Palynology 65: 47-55. Martin, Helene A. (1990b). The palynology of the Namba Formation in the Wooltana-1 bore, Callabonna Basin (Lake Frome), South Australia, and its relevance to Miocene grasslands in central Australia. Alcheringa 14: 247-255. Martin, Helene A. (1991). Tertiary stratigraphic palynology and palaeoclimate of the inland river systems in New South Wales. Geological Society of Australia Special Publication 18: 181-194. Martin, Helene A. (1992). The Tertiary of south-eastern Australia: Was it tropical? Palaeobotanist 39: 270-280. Martin, Helene A. (1993). The palaeovegetation of the Murray Basin, Late Eocene to mid Miocene. Australian Systematic Botany 6: 491-531. Martin, Helene A. (1994). Australian Tertiary phytogeography: evidence from pollen. Pp. 104-142. In: History of the Australian Vegetation: Cretaceous to Recent. (Ed. R. S. Hill) Cambridge University Press, Cambridge. Martin, Helene A. (1997). The use of ecological tolerance for the reconstruction of Tertiary palaeoclimates. Australian Journal of Botany 45: 475-492. Martin, Helene A. (1998a). Tertiary climate evolution and the development of acidity in Australia. Proceedings of the Linnean Society of New South Wales 119: 115-136. Martin, Helene A. (1998b). Late Cretaceous-Caenozoic palynology of the Poonarunna No. 1 Well, Central Australia. Transactions of the Royal Society of South Australia 122: 89-138. Martin, Helene A. (2000). Re-assignment of affinities of the fossil pollen type Tricolpites trilobatus Mildenhall & Pocknall to Wilsonia (Convolvulaceae) and a reassessment of ecological interpretations. Review of Palaeobotany and Palynology 111: 237-251. Martin, Helene A. (2006). Cainozoic climatic change and the development of the arid vegetation in Australia. Journal of Arid Enviroments. 66: 533-563. Martin, Helene A. & Gadek, P. A. (1988). Identification of Eucalyptus spathulata pollen and its presence in the fossil record. Memoirs of the Association of Australia Palaeontology 5: 311-327. Martin, Helene A. & McMinn, A. (1993). Palynology of sites 815 and 823: The Neogene vegetation history of coastal north-eastern Australia. Pp. 115-125. In: Proceedings of the Ocean Drilling Program. Scientific Results. Vol. 133. (Ed. J. A. McKenzie, P. J. Davies, & A. Palmer-Julson). Martin, Helene A. & McMinn, A. (1994). Late Cainozoic vegetation history of north-western Australia from palynology of a deep sea core (ODP site 765). Australia Journal of Botany 42: 95-102. Martin, Helene A. and Specht, R. L. (1962). Are mesic communities less drought resistant? A study on moisture relationships in dry sclerophyll forest at Inglewood, South Australia. Australian Journal of Botany 10: 106-118. Martin, Helene A. and Specht, R. L. (2005). Sclerophyll (heathy) understoreys in the Mount Lofty Ranges, South Australia. Transactions of the Royal Society of South Australia 129: 14-24. Martin, Helene A., Worral, L. & Chalson, J. (1987). The first occurrence of the Palaeocene Lygistepollenites balmei Zone in the Eastern Highlands region, New South Wales. Australian Journal of Earth Sciences 34: 359-365.

3.6.4 Quaternary palynology and vegetation

In north-eastern Queensland, the sea-level fell during the two major Ice Ages — thus exposing the Great Barrier Reef (Hopkins et al. 1996; Specht and Specht 1999a, p. 87). During each Ice Age, the tropical rainforest was confined to well-watered ‘oases’ (Kershaw 1981; Hopkins et al. 1993, 1996) while eucalypt forests invaded their old habitat (Kershaw 1981; Hopkins et al. 1993, 1996). When temperatures improved after each Ice Age, the rainforest flora re-occupied much of the landscape that had been occupied by eucalypt forests during the Ice Ages. The new species that had evolved in these ‘oases’ expanded to form the present rainforests of the region (Stocker 1981, 1988: Stocker et al. 1985). — creating a high Gamma Biodiversity in the region, with each rainforest stand of similar Alpha Biodiversity.

35 References Hopkins, M. S., Ash, J., Graham, A. W., Head, J. & Hewett, R. K. (1993). Charcoal evidence of the spatial extent of the Eucalyptus woodland expansions and rainforest contractions in Northern Queensland during the late Pleistocene. Journal of Biogeography 20: 357-372. Hopkins, M. S., Head, J., Ash, J. E., Hewett, R. K. & Graham, A. W. (1996). Evidence of a Holocene and continuing recent expansion of lowland rainforest in humid, tropical North Queensland. Journal of Biogeography 23: 737-745. Kershaw, A. P. (1974). A long continuous pollen sequence from north-eastern Queensland, Australia. Palaeogeography, Palaeoclimatology, Palaeoecology 109: 399-412. Kershaw, A. P. (1981). Quaternary vegetation and environments. Pp. 81-101. In: Ecological Biogeography of Australia. (Ed. A. Keast) Junk, The Hague. Kershaw, A. P. (1985). An extended Late Quaternary vegetation record from north-eastern Queensland and its implications for the seasonal tropics of Australia. Proceedings of the Ecological Society of Australia 13: 179-189. Kershaw, A. P. (1986). Climatic change and Aboriginal burning in north-east Australia during the last two glacial/interglacial cycles. Nature 322: 47-49. Kershaw, A. P. (1992). The development of rainforest-savanna boundaries in tropical Australia. Pp. 255-271. In: Nature and Dynamics of Forest-Savanna Boundaries. (Eds P. A. Furley, J. Proctor & J. A. Ratter) Chapman & Hall, London. Kershaw, A. P. & Nix, H. A. (1988). Quantitative palaeoclimate estimates from pollen data using bioclimatic profiles of extant data. Journal of Biogeography 15: 598-602. Kershaw, A. P. & Sluiter, I. R. (1982). Late Cenozoic pollen spectra from the Atherton Tableland, north-eastern Queensland. Australian Journal of Botany 30: 279-295. Kershaw, A. P., McKenzie, G. M. & McMinn, A. (1993). A Quaternary vegetation history of north-east Queensland from pollen analysis of ODP site 820. Pp. 107-114. In: Proceedings of the Ocean Drilling Program, Scientific Results. Vol. 133. (Eds J. A. McKenzie, P. J. Davies & A. Palmer-Julson) Kershaw, A. P. (1994). Pleistocene vegetation of the humid tropics of north-eastern Queensland, Australia. Palaeogeography, Palaeoclimatology, Palaeoecology 109: 399-412. Kershaw, A. P., Martin, Helene, A. & McEwen Mason, J. C. (1994). The Neogene: a time in transition. Pp. 299-327. In: History of Australian Vegetation: Cretaceous to Recent. (Ed. R. S. Hill) Cambridge University Press, Cambridge. Moss, P. T. & Kershaw, A. P. (2000). The last glacial cycle from the humid tropics of north-eastern Australia: Comparison of a terrestrial and marine record. Palaeogeography, Palaeoclimatology, Palaeoecology 155: 155- 176. Stocker, G. C. (1981). Regeneration of a north Queensland rainforest following felling and burning. Biotropics 13: 86- 92. Stocker, G. C. (1988). Tree species diversity in rainforests — establishment and maintenance. Proceedings of the Ecological Society of Australia 15: 39-47. Stocker, G. C., Unwin, G. L. and West, P. W. (1985). Measures of richness, evenness and diversity in tropical rainforest. Australian Journal of Botany 33: 131-137.

3.7 Rare and Endangered Australian Plants

Ray Specht and his co-workers had almost completed the IBP Conservation Survey of Australia (Specht et al. 1974) when Dr Ron Melville, recently retired from Kew Herbarium, requested them to compile a Red Book of Endangered Plant Species in Australia for the International Union for the Conservation of Nature and Natural Resources (Fisher et al. 1969).

The conservation categories of major concern were: 1. Probably extinct 3. Endangered (only small colonies remain, under adverse conditions) 4. Rare (population of adequate size, but needs constant monitoring) 5. Depleted (population originally widespread but now reduced in size; needs constant monitoring) 6. Species known only from original collection (and more information needed on its distribution) 7. Species of geographical importance (with a disjunct or isolated distribution)

With the assistance of Herbarium in each State, they rapidly compiled a list of Relict Species in the Australian Flora and their conservation (Specht et al. 1974, pp. 606-628), also identifying: —

 Relict Species that have retained floral characters generally accepted by taxonomists and morphologists as primitive.  Relict Species that have retained primitive morphological characters.

36 Over the next 20 years, J. D. Briggs and J. H. Leigh in the CSIRO Division of Plant Industry, revised and extended the list of Rare or Threatened Australian Plants (Briggs & Leigh 1988, 1996).

References Briggs, J. D. & Leigh, J. H. (1988). Rare or Threatened Australian Plants. Revised Edition. Australian National Parks and Wildlife Service Special Publication No. 14. Briggs, J. D. & Leigh, J. H. (1996). Rare or Threatened Australian Plants. Fourth Edition. CSIRO Publishing, Melbourne. Fisher, J., Simon, N. & Vincent, J. (1969). The Red Book: Wildlife in Danger. Collins, London. Melville, R. (1973). Relict plants in the Australian flora and their conservation. In: Nature Conservation in the Pacific. (Eds A. B. Costin & R. H. Groves) Australian National University Press, Canberra. Specht, R. L. et al. (1974). Primitive seed plants in the Australian flora. Pp. 606-628. In: Conservation of Major Plant Communities in Australia and Papua New Guinea. (Eds R. L. Specht, Ethel M. Roe & Valerie H. Boughton) Australian Journal of Botany Supplementary Series No. 7.

3.8 Selection of Conservation Reserves

Shortly after the publication of the assessment of the conservation of the subjectively-defined plant communities in all States and Territories of Australia (Specht et al. 1974), the Australian Heritage Commission encouraged Ray Specht to define plant communities objectively. The Botany Department of the University of Queensland documented the plant lists on all ecological surveys on the large PD10 computer of the University of Queensland (Specht A. et al. 2018), to be analysed by Mike Dale, using the classification program TWINSPAN (Hill 1973; Hill and Gauch 1980), on the large computer in the CSIRO Division of Computing Research

World Wildlife Fund (Aust.) then encouraged the Botany Department of the University of Queensland, together with the Queensland National Parks and Wildlife Service, to develop a statistical method for the selection of the most ideal location for a conservation reserve with maximum biodiversity (Bolton and Specht 1983; Purdie 1987).

References Bolton, M. P. & Specht, R. L. (1983). A Method for Selecting Nature Conservation Reserves. Australian National Parks and Wildlife Service Occasional Paper No. 8, Canberra. Hill, M. O. (1973). Reciprocal averaging: an eigen vector method of ordination. Journal of Ecology 61: 237-249. Hill, M. O. and Gauch, H. G. (1980). Detrended correspondence analyis: An improved ordination technique. Vegetation 42: 47-58. Purdie, Rosemary W. (1987). Selection of key area networks for regional nature conservation. The revised Bolton and Specht Method. Proceedings of the Royal Society of Queensland 98: 59-71. Specht, Alison, Bolton, M. P., Kingsford, B., Specht, R. L. and Belbin, L. (2018). A Story of Data Won, Data Lost and Data Re-found: The Realities of Ecological Data Preservation. Biodiversity Data Journal 6: e28073. Specht, R. L., Roe, Ethel, M. and Boughton, Valerie H. (Eds) (1974). Conservation of Major Plant Communities in Australia and Papua New Guinea. Australian Journal of Botany Supplement No. 7, Melbourne. Specht, R. L., Specht, Alison, Whelan, M. B. and Hegarty, Elwyn E. (1995). Conservation Atlas of Plant Communities in Australia. Centre for Coastal Management, Southern Cross University, Lismore, N.S.W.

3.9 Conservation by cultivation

Shortly after the publication of ‘Conservation Survey of Major Plant Communities in Australia and Papua New Guinea’ (Specht et al. 1974; Specht 1975), Ray Specht was asked to present keynote talks to several societies concerned with the ‘cultivation of native plants’ (Specht 1977, 1978, July 1978). “Conservation by Cultivation”, a slogan promoted by the Royal Botanic Gardens, Kew, UK, was adopted to ensure the survival of the multitude of ‘rare and endangered plant species’, both in Australia and internationally. This initiative comprised three major activities:

37 (1). Conservation by cultivation on the part of commercial and amateur horticulturalists, who often wished to develop even more spectacular blossoms. (2). Development of Regional Botanical Gardens for the cultivation of ‘rare and endangered’ plants of the locality. e.g. Alice Springs Botanical Garden. (3). Establishment of a ‘seed-bank’ of rare and endangered plant species to ensure the genetic material was conserved for posterity.

References Specht, R. L., Roe, Ethel M. and Boughton, Valerie H. (Eds) (1974). Conservation of Major Plant Communities in Australia and Papua New Guinea. Australian Journal of Botany Suppl. No. 7, 667 pp. Specht, R. L. (1975). A national system of ecological reserves in Australia. The report and its recommendations. Pp. 11- 21 In: A National System of Ecological Reserves in Australia. (Ed. F. Fenner) Australian Academy of Science Report No. 19. Specht, R. L. (1977). Conservation by cultivation. Pp. 3-5. In: Growing Australian Native Plants. (Ed. R. Prater) Department of Continuing Education (Namoi), University of New England, Tamworth, N.S.W. Specht, R. L. (1978). Conditions for cultivation of Australian plants. Australian Plants 9: 312-313. Specht, R. L. (July 1978). In Wildness is the Preservation of the World. Sixth Romeo Watkins Lahey Memorial Lecture, National Parks Association of Queensland, Brisbane, 12pp.

4. Syntheses

Throughout his academic career, Ray Specht was involved closely in cooperative ventures to gather, assimilate and synthesise information on ecosystems. He participated in what has been described as “the last great expedition in Australia” and continued to collect information and develop comprehensive analyses of many ecosystems.

4.1. Ethnobotany In 1948, at the age of 23 years, ecologist Ray Specht joined a group of senior scientists on the American- Australian Scientific Expedition to Arnhem Land, Northern Territory. The expedition studied the ethnology, anthropology and archaeology, in addition to the health and nutrition, of the indigenous Aborigines in relation to the marine, freshwater and terrestrial ecosystems of the region.

One of his tasks was to record the plants used by the Aborigines for food, implements/weapons, material culture, and medicine. One hundred species were recorded, together with their Aboriginal names in two languages (Specht 1958c). Over the next decades, this study initiated several in-depth investigations in Arnhem Land (Levitt 1981 in Groote Eylandt; Altman 1981 at Maningrida; Smyth and von Sturmer 1981 at Oenpelli; Russell-Smith 1985 in Kakadu; Wightman and Smith 1989 at Milingimbi; Yunupingu et al. 1995 at Yirrkala). Alan and Joan Cribb of the Botany Department of the University of Queensland were encouraged to compile ‘Wild Foods in Australia’ (Cribb and Cribb 1974) and ‘Wild Medicine in Australia’ (Cribb and Cribb 1981). Beth Gott (1982) compiled an article on ‘Ecology and Root Use by the Aborigines in Southern Australia’.

The seasonal cycle of food-gathering by hunter-gatherers that was recorded in Arnhem Land has been explored by other scientists — in Cape York Peninsula, Central Australia and South Australia (Specht and Specht 1999a, pp. 143-149).

38 The mammologist, Dave Johnson, noted that the Aborigines of Arnhem Land fired the savanna understorey on a two- to three-year cycle — to ensure that the tall spear-grasses (Shaw 1957) did not suppress the smaller grasses in which the populations of small mammals could regenerate (Johnson 1961). These observations were confirmed in the controlled experiments conducted by Dick Braithwaite of CSIRO at Kapalga in the Northern Territory (Braithwaite 1966).

The disposal of food-wastes around camp-sites was noted by Margaret McArthur (1960, p. 123). Waste products were simply thrown away into the native vegetation where, if not eaten by the camp-dogs, decomposed — thus increasing the fertility of the nutrient-poor soils. Defaecation and urination added further nutrients to soils close to camps, changing the ecological conditions in favour of nutrient- dependent (often weed) species. Similarly, near a traditional aboriginal camp ground in north-eastern New South Wales, blady grass (Imperata cylindrica) has invaded the heathy understorey (Specht, Alison unpublished, Specht and Specht 1999a, pp. 154-155). Later, Hynes and Chase (1982) reported that seeds of edible rainforest fruits that were discarded around camps on Cape York Peninsula had germinated, giving rise to the phenomenon they described as domiculture.

References Altman,J. C. (1981, unpublished). An inventory of flora and fauna utilised by eastern Gunwinggu in the Mann/Liverpool region, north central Arnhem Land. Australian Institute of Aboriginal Studies, Canberra. Braithwaite, R. W. (1996). A healthy savanna, endangered mammals and Aboriginal burning. Pp. 91-102. In: Country in Flames. (Ed. D. B. Rose). Australian National University, North Australia Research Unit, Biodiversity Series, Paper No. 3, Darwin. Cawte, J. (1996). Healers of Arnhem Land. University of New South Wales Press, Sydney. Cribb, A. B. & Cribb, Joan W. (1974, 1990) Wild Food in Australia. Collins, Sydney. Cribb, A. B. & Cribb, Joan W. (1981, 1990) Wild Medicine in Australia. Collins, Sydney. Gott, Beth (1982). Ecology and root use by the Aborigines of southern Australia. Archaeology in Oceania 17: 59-67. Hynes, R. A. & Chase, A. K. (1982). Plants, sites and domiculture: Aboriginal influence upon plant communities in Cape York Peninsula. Archaeology in Oceania 17: 38-50. Johnson, D. H. (1961). Mammals of the Arnhem Land Expedition. Pp. 427-515. In: Records of the American- Australian Expedition to Arnhem Land. Vol. 4. Zoology. (Ed. R. L. Specht). Melbourne University Press, Melbourne. Levitt, Dulcie (1981). Plants and People. Aboriginal Uses of Plants on Groote Eylandt. Australian Institute of Aboriginal Studies, Canberra. McArthur, Margaret, McCarthy, F. D. & Specht, R. L. (2000). Nutrition studies (1948) of nomadic Aborigines in Arnhem Land, northern Australia. Asia Pacific Journal of Clinical Nutrition 9: 215-223. McCarthy, F. D. & McArthur, Margaret (1960). The food quest and the time factor in Aboriginal economic life. Pp. 145-194. In: Records of the American-Australian Expedition to Arnhem Land. Vol. 2. Anthropology and Nutrition. (Ed. C. P. Mountford). Melbourne University Press, Melbourne. Russell-Smith, J. (1985). Studies in the Jungle People, fire and monsoon forest. Archaeological Research in Kakadu National Park. Australian National Parks and Wildlife Service, Canberra. Special Publication 13: 241-267. Shaw, N. H. (1957). Bunch spear grass dominance in burnt pastures in south-eastern Queensland. Australian Journal of Agricultural Research 8: 325-334. Smyth, D. and von Sturmer, J. (1981, unpublished). The use of plants by the Aboriginal people in the Oenpelli region of western Arnhem Land. Australian Institute of Aboriginal Studies, Canberra. Specht, R. L. (1958). An introduction to the ethno-botany of Arnhem Land. Pp. 479-504. In: Records of the American-Australian Expedition to Arnhem Land. Vol. 3. Botany and Plant Ecology. (Eds R. L. Specht & C. P. Mountford). Melbourne University Press, Melbourne. Specht, R.L. (1990). Changes in the eucalypt forests of Australia as a result of human disturbance. Pp. 177-197. In: The Earth in Transition. Patterns and Processes of Biotic Impoverishment. (Ed. G. M. Woodwell). Cambridge University Press, New York. Specht, R. L. (2012). American-Australian Scientific Expedition to Arnhem Land (1948) — Its Long-Range Impact. The Open Ecology Journal 2012, Vol. 5: 30 pages. Specht, R. L. & Specht, Alison (1999, 2002). Aboriginal impact. Pp. 132-156. In: Australian Plant Communities. Dynamics of Structure, Growth and Biodiversity. Oxford University Press, Melbourne. Tindale, N. B. (1981). Desert aborigines and the southern coastal people: Some comparisons. Pp. 1853-1884. In: Ecological Biogeography of Australia. (Ed. A. Keast). Junk, The Hague. Wightman, G. M. and Smith, N. M. (1989). Ethnobotany, Vegetation and Floristics of Milingimbi, Northern Australia. Northern Territory Botanical Bulletin no. 6, Daewin.

39 Yunuping, B., Yunuping-Marika, L., Marika, D., Marika, B., Marika, R. & Wightman, G. (1995). Rirratjingu Ethnobotant: Aboriginal Plant Use from Yirrkala, Arnhem Land, Australia. Northern Territory Botanical Bulletin No. 21.

4.2. American-Australian Scientific Research Expedition to Arnhem Land (1948)

As one of the Australian team on the National Geographic Expedition to Arnhem Land in 1948, Ray Specht helped organise the 25th and 50th Reunions in 1973 and 1998.

For the 25th Reunion in 1973, he collated the ‘camp-fire’ songs that the members had composed on the Expedition.

For the 50th Reunion in 1998, he collated the ‘Personal Experience on the Coast of Adventure’ of Expedition members. Dorita Thomson recalled her husband, Donald Thomson, who had helped to calm the Aborigines after the spearing of Constable McColl on Woodah Island. Velma Leeding and Carol Lenfer wrote tributes to the Aborigines at Umbakwumba on Groote Eylandt. Max Finlayson wrote an account of Aboriginal management of the Blyth/Liverpool River System.

During May 2006, Martin Thomas and Sally May spent three days interviewing Ray Specht on behalf of the National Library of Australia; later, during March 2009, Marie-Louise Ayres of the National Library visited him to obtain further information.

Margo Neales of the National Museum of Australia invited contributors for an international symposium, ‘Barks, Birds and Billabongs’, to celebrate the 60th anniversary of the Arnhem Land Expedition in 2009. The contributions were published in ‘Exploring the Legacy of the 1948 Arnhem Land Expedition’ by the Australian National University E-Press (Thomas and Neale 2011). Ray Specht summarised the long-range impact of the Arnhem Land Expedition in an article in The Open Ecology Journal (Specht 2012).

For the 70th Anniversary, John Pickrell (2018) recalled ’The Last Great Expedition’ in the January-February Edition of the Australian Geographic.

References Neale, Margo (2008). Barks, Birds and Billabongs. Exploring the Legacy of the 1948 American-Australian Scientific Expedition to Arnhem Land. International Symposium. National Museum of Australia. 16-20 November 2009. National Museum of Australia, Canberra, 108 pp. McArthur, Margaret, McCarthy F. D. & Specht, R. L. (2000). Nutritional studies (1948) of nomadic Aborigines in Arnhem Land, northern Australia. Asia Pacific Journal of Clinical Nutrition 9: 215-223. Pickrell, J. (2018). 70th Anniversary - ‘The Last Great Expedition’. Australian Geographic Jan.-Feb. 2018; 94-103. Specht, R. L. (1973). ’Song Book’. Arnhem Land Reunion, Sydney. Specht, R. L. (2012). American-Australian Scientific Expedition to Arnhem Land (1948): Its Long-Range Impact. The Open Ecology Journal 5: 53-83. Specht, R. L. & Specht, Alison, (1998). ‘Personal Experience on the Coast of Adventure’ Arnhem Land Reunion, Sydney. Southern Cross University, Lismore, N.S.W. Thomas, M. & Neale, Margo (Eds) (2011). Exploring the Legacy of the 1948 Arnhem Land Expedition. Australian National University E-Press, Canberra, 471 pp.

4.3. Ecosystems of the World (Elsevier, Amsterdam) While he was visiting Melbourne during 1965, Dr David W. Goodall was invited to attend a Planning Committee meeting for the International Biological Program (IBP), Section on Ecosystem Productivity.

40 He later became Head of the IBP Arid Zone Biome Program, based in Utah in the United States. During that part of his career, he was approached by Elsevier Publishers to become General Editor for a 30 Volume study of the Ecosystems of the World — a massive task that took the next 40 years.

Goodall invited Prof. Val Chapman of Auckland University, New Zealand, to edit Volume 1. Wet Coastal Formations (Mangroves and Salt Marshes). Chapman sought the advice of algologist, Dr Sophie Ducker of Melbourne University, who referred him to Ray Specht in the University of Queensland (with Arnhem Land, South Australian and Victorian experience). Peter Saenger (Sophie’s former algology student who was in Queensland studying mangrove reclamation in Gladstone Harbour) and Marion Specht (an animal ecologist who had studied the mangrove fauna of Moreton Bay) formed the team to prepare an article on’ Mangal and coastal salt marsh communities in Australasia.’ (Saenger et al. 1977).

Shortly afterwards, Goodall invited Ray Specht to edit the volume on Mediterranean-Type Shrublands. As he was now in subtropical Queensland, Specht declined; so Goodall suggested that he edit the volume on Heathlands and Related Shrublands, that grew on nutrient-poor soils throughout the World. There were so many contributions that the contributions were split into two volumes — Descriptive Studies and Analytical Studies. As nutrient-poor soils — on sandstones, quartzites, lateritic podsols and sand-plains — are widespread throughout the continent of Australia, many Australian scientists were invited to contribute.

Contributors (Botany): H. T. Clifford (pollination & seed dispersal), R. G. Coleman (polyphosphate metabolism), D. J. Connor (water relations), D. Doley (water relations), Wendy E. Drake (pollination & seed dispersal), A. S. George, A. M. (W.A. heathlands), Gill (fire-tolerance), A. R. Glenn (Phytophthora cinnamomi), R. H. Groves (fire-tolerance), A. J. M. Hopkins (seasonal rhythms in W.A. heathlands), B. B. Lamont (specialized roots), N. Malajcuk, (specialized roots & Phytophthora cinnamomi), N. G. Marchant (W.A. heathlands), P. G. Ozanne (phosphorus toxicity), R. F. Parsons (effect of salt-spray), R. W. Rogers (flowering rhythms), R. L. Specht (water relations, seasonal rhythms, ployphosphate & conservation), J. S. Womersley (heathlands in Malesia) and D. J. Yates (optical properties).

Contributors (Zoology): R. Y. Berg (ants), P. C. Catling (fauna of SE Australian heathlands), P. D. Dwyer (vertebrate fauna), S. J. Edmonds (invertebrate faunal rhythms in overstorey), G. J. Ingram (vertebrate fauna), J. Kikkawa (vertebrate fauna), A. R. Main (fire-tolerance of animals), A. E. Newsome (fauna of SE Australian heathlands), Marion M. Specht (invertebrate faunal rhythms in litter).

As Francesco di Castri, the Editor of Mediterranean-Type Shrublands, had accepted a position as ecologist with UNESCO in Paris, Ray Specht was co-opted to help edit this volume. It was discovered that Francesco believed that the vegetation of southern Australia and Cape Province in South Africa was all Heathlands and Related Shrublands, all on nutrient-poor soils, which had been covered in Volume 9. Fortunately, Prof. Eugene Moll of Cape Town University and Ray Specht of the University of Queensland quickly found authors with expertise on the ecology of the tall shrubland vegetation (termed mallee in Australia) on richer soils — many areas of which had been cleared for agriculture.

Later, Ray was invited to write chapters on Wetland Forests and Dry Coastal Ecosystems in Australia.

Ray Specht, therefore, participated in the production of five of these Volumes —Wet Coastal Formations (1977), Heathlands and Related Shrublands (1979, 1981), Mediterranean-Type Shrublands (1981), Wetland Forests (1990) and Dry Coastal Ecosystems (1993, 1997).

References Di Castri, F., Goodall, D. W. and Specht, R. L. (Eds) (1981). Ecosystems of the World. Vol. 11. Mediterranean- Type Shrublands. Elsevier, Amsterdam.

41 Saenger, P., Specht, Marion M., Specht, R. L. and Chapman, V. J. (1977). Mangal and coastal salt marsh communities in Australasia. Pp. 293-345. In: Ecosystems of the World. Vol. 1. Wet Coastal Formations. (Ed. V. J. Chapman) Elsevier, Amsterdam. Specht, R. L. (Ed.) (1979). Ecosystems of the World. Vol. 9A. Heathlands and Related Shrublands. Descriptive Studies. Elsevier, Amsterdam. Specht, R. L. (Ed.) (1981). Ecosystems of the World. Vol. 9B. Heathlands and Related Shrublands. Analytical Studies. Elsevier, Amsterdam. Specht, R. L. (1990). Forested wetlands in Australia. Pp. 387-406. In: Ecosystems of the World. Vol. 15. Wetland Forests (Eds A. E. Lugo, M. M. Brinson & Sandra Brown) Elsevier, Amstrdam. Specht, R. L. (1993). Dry coastal ecosystems of Australia — An overview of the dune vegetation. Pp. 223- 237. In: Ecosystems of the World. Vol. 2B. Dry Coastal Ecosystems: Africa, America, Asia and Oceania (Ed. E. van der Maarel) Elsevier, Amsterdam. Specht, R. L. (1997). Ecosystem dynamics in coastal dunes in eastern Australia, Pp. 483-495. In: Ecosystems of the World. Vol. 2C. General Aspects. (Ed. E. van der Maarel) Elsevier, Amsterdam.

4.4. Ecological Biogeography of Australia (Dr W. Junk, The Hague)

In 1956, the Editor of Dr W. Junk’s Monographiae Biologicae series visited Australia and encouraged Prof. R. L. Crocker in the Botany Department, University of Sydney, and C. S. Christian, CSIRO Division of Land Research and Regional Survey, to be editors of the first volume in their new series, ‘Biogeography and Ecology’ in Australia (1959). The volume was so successful that Junk invited Alan Keast, who was a Visiting Professor at Griffith University, to edit ‘Ecological Biogeography of Australia’ (1981). Alan invited Ray Specht to co-ordinate the contributions for Part two: ‘The Flora of Australia’ for this volume.

Chapter 8: Specht, R. L. (Major vegetation formations in Australia), Chapter 9: Specht, R. L. (Eco-phyiological principles), Chapter 10: Beard, J. S. (Phytogeographic regions). Chapter11: Dettmann, Mary E. (Cretaceous flora), Chapter12: Christophel, D. C. (Tertiary mega-fossils flora), Chapter13: Martin, Helene A. (Tertiary flora), Chapter14: Beadle, N. C. W. (Angiosperm flora). Chapter15: Johnson, L. A. S. & Briggs, Barbara G. (Three old southern families – Myrtaceae, Proteaceae & Restionaceae), Chapter16: Page, C. T. & Clifford, H. T. (Australian conifers and ferns), Chapter17: Pryor, L. D. & Johnson, L. A. S. (Eucalyptus), Chapter18: Clifford, H. T. & Simon, B. K. (Australian grasses), Chapter19: Barlow, B. A. (Australian mistletoes), Chapter20: Specht, R. L. (Biogeography of halophytes), Chapter21: Rogers, R. W. & Stevens, G. N. (Lichens), Chapter22: Webb, L. J. & Tracey, J. G. (Australian rainforests), Chapter23: Beadle, N. C. W. (Vegetation of the Arid Zone), Chapter24: Costin, A. B. (Vegetation of high mountains), Chapter25: Charles-Nelson, E. (Genetic systems & phytogeography of southern Australia), Chapter26: James, S. H. (Cyto-evolutionary patterns), Chapter27: Specht, R. L. (Evolution of the Australian flora — Some generalizations).

4.5. Mediterranean-type ecosystems

When the International Biological Program was launched in 1965, the Unites States Academy of Science decided to combine the three sections — productivity, production processes and conservation (Specht 1966) — for the study of terrestrial ecosystems under Biome Studies.

Hal Mooney of Stanford University, California, launched Mediterranean-type Biome Studies with Franceso di Castri of the Catholic University of San Diego, Chile at a conference in 1972. Ray Specht’s paper,

42 presented in absentia (Specht 1973) inspired MEDECOS (Mediterranean-Type Ecosystem Studies). Phil and Patsy Miller with Joachim Kummerow of SERG, San Diego, California, spent Christmas camped in Dark Island heathland to study our research. Many nations subscribed for copies of the Australian Journal of Botany to study our articles. Ecologists from every Mediterranean-Type Climate nations then met every two years to discuss research on ecosystems in their country.

References Specht, R. L. (1966). The International Biological Program (I.B.P.), Section PCT: Productivity, production processes and conservation of terrestrial communities. Australian Journal of Science 28: 377-380. Specht, R. L. (1973). Structure and functional response of ecosystems in the Mediterranean climate of Australia. Pp. 113-120. In: Mediterranean Type Ecosystems. Origin and Structure. (Eds F. di Castri & H. A. Mooney) Springer- Verlag, Berlin.

To co-ordinate the diverse contributions from each nation, it was proposed to collate the data on: —

(1) Climatic controls on growth and community distribution;

(2) Regional variations in plant nutrient status and the effects on sclerophylly;

(3) Species richness of plants, vertebrates and soil fauna as a function of stand architecture, climate, soil moisture and substrate stability;

(4) Ecomorphological characters and seasonality — the effect of changes in these parameters along environmental gradients on community function.

The results of these studies were collated in Mediterranean-type Ecosystems. A Data Source Book. (Specht 1988).

Chapters contributed by Ray Specht and associates:—

1. Arianoutsou, Margareta & Paraskevopoulos S. (with Hutley, L. B. & Specht, R. L.) (1988).Climatic and

Vegetation Data for Greece. (pp. 125- 126). 2 Berliner, Ruhama, R. L. Specht & Hutley, L. B. (1988). Climatic and Vegetation Data for Israel. (pp. 126- 128). 3 Catarino, F. M. & Correia A. I. D. (1988). Climatic and Vegetation Data for Portugal (pp. 129-130). 4 Gracia, C. A. (1988). Climatic and Vegetation Data for East Spain. (pp. 131-133). 5 Hajek, E. J., Gajardo, R. & Molina, J. D. with Specht, R. L. & Hutley, L. B. (1988). Climatic and Vegetation Data for Chile. (pp. 113-116). 6 Specht, R. L. (1988). Climatic and Vegetation Data for Arizona, U.S.A. (pp. 110-112).

7 Specht, R. L. & Moll. E. J. (1988). Climatic and Vegetation Data for Cape Province, South Africa. (pp. 135- 136). 8 Zedler, P. H. & Specht, R. L. (1988). Climatic and Vegetation Data for California, U.S.A. (pp. 106- 110).

As the Mediterranean-type vegetations (and other communities) of South Africa and Chile in South America are related to the Australian vegetation — being formerly closely united on Gondwanaland — a detailed analysis was, later, made of both these countries (Specht and Moll 2014; Specht et al. 2015).

References Specht, R. L. (Ed.) (1988). Mediterranean-type Ecosystems. A Data Source Book. Kluwer Academic Publishers, Dordrecht. Specht, R. L. and Moll, E. J. (2014). Structure and alpha biodiversity of major plant communities in South Africa, a close biogeographical relation with Australia. Macrothink Institute Journal of Environment & Ecology 5: 48-88.

43 Specht, R. L., Montenegro, Gloria and Dettmann, Mary E. (2015). Structure and alpha biodiversity of major plant communities in Chile, a distant biogeographical relation with Australia. Macrothink Institute Journal of Environment & Ecology 6: 21-47.

4.6. ‘Australian Plant Communities. Dynamics of Structure, Growth and Biodiversity’ (Specht, R. L. & Specht, Alison (1999, 2002). Oxford University Press, Melbourne).

This major work drew together a lifetime of endeavour to understand the fundamentals of the structure and functioning of Australian plant communities. In keeping with the basic IBP program, it was presented in two parts, with the following chapter details. Part 1. Australian plant communities: Description Chapter 1. Energy/biomass system; Chapter 2. Dynamics; Chapter 3. Structural classification; Chapter 4. Floristic classification; Chapter 5. Australian plant communities during the Late Cretaceous; Chapter 6. Australian plant communities during the Cainozoic; Chapter 7. Australian plant communities today; Chapter 8. Aboriginal impact

Part 2. Australian Plant Communities: Community Physiology Chapter 9. Energetics; Chapter 10. Temperature; Chapter 11. Evaporative aerodynamics; Chapter 12.Available water; Chapter 13. Eco-physiological leaf attributes; Chapter 14. Waterlogging; Chapter 15. Nutrient deficiencies; Chapter 16. Nutrient toxicities; Chapter 17. Biodiversity and energetics; Chapter 18. Monitoring; Chapter 19. Scientific management

References Specht, R. L. & Specht, Alison (1999, 2002). Australian Plant Communities. Dynamics of Structure, Growth and Biodiversity. Oxford University Press, Melbourne. Specht, R. L. and Specht, Alison (2013). Australia: Biodiversity of ecosystems. Pp. 291-306. In Encyclopedia of Biodiversity. (Ed. S. Levin) Elsevier, Amsterdam. Republished in Reference Module in Life Sciences, Elsevier 2016.

5. Education Ray Specht spent his entire career as an academic, successively in the Departments of Botany in the Universities of Adelaide (Lecturer and Senior Lecturer), Melbourne (Reader) and Queensland (Professor and Head of Department). In addition to his assigned responsibilities in the field of ecology at university level, he took a leading role in the promotion of biological education in schools and in the general community.

A. Teaching material for Queensland tertiary students in botany

Clifford, H. T. (1981). Key to Families of Flowering Plants. Pp. 113-168. In: Flora of Australia. (Ed. A. S. George) Australian Government Publishing Service, Canberra. Clifford, H. T. (1972). Eucalypts of the Brisbane Region. Queensland Museum Publication, Brisbane. Clifford, H. T. & Constantine, Jean (1980) Ferns, Fern Allies and Conifers of Australia. University of Queensland Press, St Lucia. Clifford, H. T. & Ludlow, Gwen (1978) Keys to the Families and Genera of Queensland Flowering Plants (Magnoliophyta). University of Queensland Press St Lucia.

44 Clifford, H. T. & Specht, R. L. (1979) The Vegetation of North Stradbroke Island. University of Queensland Press, St Lucia. Clifford, H. T. & Stephenson, W. T. (1975). An Introduction to Numerical Classification and Taxonomy. Academic Press, New York. Clifford, H. T. & Watson, L. (1977). Identifying Grasses: data, methods and illustrations. University of Queensland Press. Cribb, A. B. (1985) Key to the Queensland Marine Algae. Unpublished, Botany Department, University of Queensland. Dahlgren, R. M. T. & Clifford, H. T. (1982). The Monocotyledons. A Comparative Study. Academic Press, New York. Dahlgren, R. M. T., Clifford, H. T. & Yeo, P. F. (1985). The Families of the Monocotyledons: Structure, Evolution and Taxonomy. Springer-Verlag, Berlin. Specht, R. L. (1972). The Vegetation of South Australia. Government Printer, Adelaide. (Pages 1-71 of this Handbook contained material for teaching, developed for computer modelling classes — see the following publication) Specht, R. L. & Specht, Alison (1999, 2002). Australian Plant Communities. Dynamics of Structure, Growth and Biodiversity. Oxford University Press, Melbourne.

B. Biological science curriculum studies – ‘The Web of Life’

Ray Specht’s education at Adelaide Teachers College and the University of Adelaide was to prepare him to develop Biological Science in Secondary Schools in South Australia. The 1948 Arnhem Land Expedition changed this direction.

After the Russians launched Sputnik in 1958, scientists in the United States began programs to improve the teaching of science in Secondary Schools — Physical Science Curriculum Studies (PSCS), Chem-Studies, Biological Science Curriculum Studies (BSCS). For each of these sciences, some 1,000 educators and 1,000 scientists collaborated. The ‘Major Ideas’ (the most up-to-date principles) were defined for teaching by the ‘Inquiry Method’ in discussion groups, practicals, etc. — not by rote learning. As the study of Biology is so vast, BSCS decided to produce three teaching courses — Study at the Cellular Level; Study at the Whole Organism Level; Study at the Ecosystem Level.

Because there are marked differences between the flora and fauna of the United States and Australia, educators in South Australia and Victoria were asked to combine these three courses into one — ‘The Web of Life’ — with the materials published by the Australian Academy of Science, after mortgaging their new building.

Ray Specht and his working group were encouraged to launch ‘The Web of Life’ in Queensland. A small group from 25 pilot schools in south-eastern Queensland became involved. David Morgan and team from Melbourne delivered a week’s training course in the Botany Department of the University of Queensland during December 1967. Over the next two years, the Botany Department held monthly ‘chats’ on any problems that a teacher may have encountered. At the end of the second year (November 1969), they held a training course for 100 teachers from all over Queensland, repeated with another 100 new teachers two years later (December 1971).

Ray’s wife, Marion, who had taught ‘The Web of Life’ at Brisbane Girls Grammar School during the pilot course, was appointed as Queensland Liaison Officer by the Australian Academy of Science for one to two days per week — a task that often extended to five or seven days per week. Once a month, she travelled north to various ‘coastal’ Secondary Schools where teachers from the ‘inland’ discussed any problems that they had with the course.

With the Queensland Education Department, the team ensured that equipment was available at the correct time for each class throughout the State. The Education Department appointed three teachers to ensure biological material was delivered on time.

45 Booklets describing the major ecosystems, flora and fauna in each region were prepared for the information of teachers new to the area.

Ray maintained the Queensland contact with the Australian Academy of Science as a member of the Committee on Biological Education, 1968-1985. The ‘Web of Life’ continued to flourish in Queensland well into the 1980s. Even today (1994), some Queensland teachers trained in the ‘Web of Life’ Teachers’ Preparation Courses in the 1960s and 1970s talk glowingly of this Australian biology course and still adhere to the Inquiry-Method of teaching (Specht and Specht, Marion 1995).

References Morgan, D. G. et al. (1967). Biological Science: The Web of Life. Australian Academy of Science, Canberra. Plus Teacher’s Guide and Student’s Manual Parts 1 & 2. Specht, R. L. and Specht, Marion M. (1995). The 'Web of Life' in Queensland. (Australian Academy of Science). Proceedings of the Royal Society of. Queensland 105: 55-60.

C. University of Queensland’s 75th celebrations

The 75th Anniversary celebrations of the opening of the University of Queensland in 1913 included a conference highlighting the distinctive attributes of the Queensland flora and the contributions of members of the Department of Botany to an understanding of this flora.

References Clifford, H. T. & Specht, R. L. (Eds) (1986). Tropical Plant Communities. Their Resilience, Functioning and Management in Northern Australia. Utah Foundation & University of Queensland, Department of Botany, St Lucia. Sanders, Dorothea F. & Specht, R. L. (1987). Thomas Harvey Johnston M. A., D. Sc. Professor of Biology, 1919-1923. Pp. 59-62. In: Vivant Professores. (Ed. Helen Gregory). University of Queensland Library, St Lucia.

6. Community Services

In addition to fundamental research work on plant communities, Ray Specht became involved in research topics related to plants and vegetation in Queensland. These research projects ranged from health effects of the seasonal growth of plants, the influence of mining and industrial environments on vegetation and the durability of wooden power poles.

6.1. Childhood Asthma — associated with seasonal growth of eucalypt foliage Shortly after Ray Specht had arrived in Brisbane, a report on the research to discover the cause of childhood asthma (in children under three years of age) was published in the ‘Courier Mail’. The numbers of little children, who had been admitted to hospital with an asthma attack, had been recorded monthly for several years; these values peaked in both autumn and spring.

Studies on the amount of tropical grass pollen, fungal spores, pollutant particles in the atmosphere were not correlated — even under the inversion layer that sometimes occurred in the Brisbane River valley during autumn. Off-shore winds, however, seemed to reduce hospital admissions.

Perchance, while reading this asthma report in the ‘Courier Mail’, Ray noticed new leaves being formed on the eucalypt (Corymbia intermedia) outside his bedroom window — during autumn, a season much later than the summer growth-rhythm in the South. He obtained a grant from the Queensland Asthma Foundation to follow the seasonal growth of eucalypts in the Brisbane region (Specht and Brouwer 1975).

46 Dr Derrick of the Queensland Institute of Medical Research continued his study of hospital admissions of childhood asthma and they confirmed that these admissions correlated with the autumn and spring foliage-growth-rhythms of eucalypts in the Brisbane area (Specht et al. 1975). Tiny insects and mites feed on the edges of developing leaves; their droppings then cloud the atmosphere (Specht and Specht 1999b, pp. 375-376).

Dr Derrick’s sister, who was a scientist in the Australian Medical Research Institute in Melbourne, obtained data on childhood asthma admissions to hospitals in Victoria; these were correlated with the summer growth-rhythm of eucalypts in the South.

Ray’s daughter, Alison, continued the study of the relationship between seasonal shoot growth in eucalypts and childhood asthma in the Brisbane region, and was awarded the first Lillian Roxon Asthma Grant that enabled her and her husband, Ian Whyte, to attend an International Asthma Conference in

Torremolinos, Spain (Specht and Whyte 1978).

References Specht, Alison and Whyte, I. M. (1978). Air spora studies in vegetation in Brisbane and its possible relationship with asthma. In: Proceedings INTERASMA, Torremolinos, Spain, 1977. Specht, R. L. and Brouwer, Yvonne M. (1975). Seasonal shoot growth of Eucalyptus spp. in the Brisbane area of Queensland (with notes on shoot growth and litter fall in other areas of Australia). Australian Journal of Botany 23: 459-474. Specht, R. L. and Specht, Alison (1999). Asthma allergens. Pp. 375-376. In: Australian Plant Communities. Dynamics of Structure, Growth and Biodiversity. Oxford University Press, Melbourne. Specht, R. L., Brouwer, Yvonne M. and Derrick, E. H. (1975). Seasonal waves of asthma: A possible botanical cause. International Journal of Biometeorology 19: 28-36.

6.2. The Mining Industry

6.2.1. Bauxite mining, Weipa The bauxite mining industry at Weipa on Cape York Peninsula, Queensland, was concerned about the environmental impact of mining on ‘rare-and-endangered species’. During 1976, Ray Specht accompanied their mining engineer, Richard Salt, on two surveys of the vegetation at Weipa — the first in mid-year, the second at the end of the Wet Season to check for any short-lived annuals. Sally Reynolds of the Queensland Herbarium identified the plants collected to check for any ‘rare-and- endangered species’ that may be on the area to be mined (Specht et al. 1977).

It was fortunate that a hydrology survey had been made to determine the depth of the underlying water table below future mining excavations. The depth of this water table gradually became lower towards the coast until it enabled a constant supply of fresh-water to bathe the deep roots of a closed- forest community (Specht et al. 1977) — thus, clearly showing that the boundary from eucalypt forest to rainforest was related to the continuous availability of water — not to bush-fires.

6.2.2. Fluoride emission from aluminium smelters

47 In 1973, Comalco Limited managers approached the Botany Department to investigate the possible effects of gaseous fluoride emissions on tropical native vegetation as part of the environmental impact statement for a proposed aluminium smelter at Gladstone, in central Queensland. This work had to be carried out in custom-built fumigation cabinets and in the laboratory, and was followed by similar studies for established smelters in New South Wales (Doley 1984, 1986a), Victoria, Tasmania and New Zealand, and for proposed smelters in Queensland (Doley 1988, 1989), Victoria and Western Australia. The accumulation of information resulted in the publication of a handbook (Doley 1986b) and in a 30- year advisory association with industry (Doley et al. 2004; Doley 2010;).

6.2.3. Sulphur dioxide emission from copper refiners During 1974, Ken Dredge and Cliff Thompson from Mt Isa Mines approached the Botany Department to study the effect of sulphur-dioxide emitted from their smelting chimney. Ray Specht and Rod Rogers examined the death of the Triodia hummock grassland to the west of the Mine, downstream from the chimney — before the stack had been increased in height. On a grant from Mt Isa Mines, Dave Doley supervised Honours student, John Henry, in a study of the effects on vegetation of short-term high- concentration sulphur dioxide exposure events.

6.2.4. Serpentinite, a nickel-bearing deposit During 1989, Roger Reeves, Massey University, Palmerston North, New Zealand, was encouraged to spend his sabbatical in the Botany Department, University of Queensland, to investigate the vegetation on serpentinite, a nickel-bearing rock, in Queensland. He and Ray Specht firstly studied the vegetation on the large deposit between Rockhampton and Marlborough in Central Queensland; George Batianoff of the Queensland Herbarium identified their collection.

The first specimen they collected was a species of Stackhousia that proved to have the world’s highest level of nickel in its leaves. George renewed its original name, Stackhousia tryoni, named by F. M. Bailey after entomologist, Henry Tryon, who had collected the species on a nickel-bearing deposit on South Percy Island. The level of nickel in the leaves of this plant far exceeded any of our collections from serpentinite deposits at Greenvale, Rockhampton-Marlborough, and Widgee Mountain in Queensland or at Baryulgil in north-east New South Wales. (Specht, Alison et al. 2001).

In the edaphic gradient from shallow to deep soils on this relatively impermeable serpentinite rock in the Rockhampton-Marlborough area, the vegetation increased in structure to an open-forest, about 25m tall. Wherever seepage flowed into deep soils, a closed-forest (rainforest) developed, fringed with eucalypts, almost 40m tall. (Batianoff and Specht 1992; Batianoff et al. 1990, 1991, 1997).

6.2.5. Mine-reclamation During Ray Specht’s second visit (March 1977) to Weipa to seek for any annual species that may have germinated during the Wet Season, Richard Salt organised the first Mine Reclamation Conference. Ray

48 Specht was then invited to give a key-note address on ‘Environmental planning and management for mining and energy’ at the Mine Rehabilitation Workshop in Darwin, June 1986.

Subsequently, Dave Doley, Dave Lamb and Dave Yates were associated with the Centre for Mined Land Rehabilitation in the Sustainable Minerals Institute of The University of Queensland, where they applied the principles of biophysics, plant physiology and plant ecology to the re-establishment of functioning ecosystems on landscapes that had been radically disturbed by mining operations, both in Australia and overseas.

References Batianoff, G. N. and Specht, R. L. (1992). Queensland (Australia) serpentine vegetation. Pp. 109 – 128. In: The Vegetation of Ultramafic (Serpentine) Soils. (Eds A. J. M. Baker, J. Proctor & R. D. Reeves) Intercept Publications, Andover, England. Batianoff, G.N., Reeves, R.D. and Specht, R.L. (1990). Stackhousia tryonii Bailey: a nickel-accumulating serpentinite- endemic species of central Queensland. Australian Journal of Botany. 38 : 121-130. Batianoff, G.N., Specht, R.L. and Reeves, R.D. (1991). The serpentinite flora of the humid subtropics of eastern Australia. Proceedings of the Royal Society of Queensland 101: 137-158. Batianoff, G. N., Reeves, R. D. and Specht, R. L. (1997). The effect of serpentine on vegetation structure, species diversity and endemism in Central Queensland. Pp. 44-50. In: The Ecology of Ultramafic and Metalliferous Areas. (Eds T. Jaffré, R. D. Reeves & T. Becquer) Centre ORSTOM de Nouméa, New Caledonia. Doley, D. (1984). Experimental analysis of fluoride susceptibility of grape vine (Vitis vinifera L.). Foliar fluoride accumulation in relation to ambient concentration and wind speed. New Phytologist, 96, 337-351. Doley, D. (1986a). Experimental analysis of fluoride susceptibility of grape vine (Vitis vinifera L.). Leaf development during four successive seasons of fumigation. New Phytologist, 103, 325-340. Doley, D. (1986b). Plant – Fluoride Relationships. Inkata Press, Melbourne. Doley, D. (1988). Fluoride-induced enhancement and inhibition of photosynthesis in four taxa of Pinus. New Phytologist 110: 21-31. Doley, D. (1989). Fluoride-induced enhancement and inhibition of shoot growth in four taxa of Pinus. New Phytologist 112: 543-552. Doley, D. (2010) Rapid quantitative assessment of visible injury to vegetation and visual amenity effects of fluoride air pollution. Environmental Monitoring and Assessment 160, 181-198. Doley, D., Hill, R.J. and Riese, R.H. (2004). Environmental fluoride in Australasia: Ecological effects, regulation and management. Clean Air and Environment 38(2), 35-55. Specht, Alison, Forth, F., and Steenbeeke, G. (2001). The effect of serpentine on vegetation structure, composition and endemism in northern New South Wales, Australia. South African Journal of Science 97: 521-529. Specht, R. L. (2007). Species richness of rainforest stands on non-serpentinite and serpentinite substrates in the Rockhampton – Marlborough area of Central Queensland. Proceedings of the Royal Society of Queensland 113: 17-35. Specht, R. L., Batianoff, G. N. and Reeves, R. D. (2005). The vegetation continuum on the serpentinite soil catena. pp. 23-40. In: The Rockhampton-Marlborough Area in Marlborough Serpentine, Landscape, Management and Conservation. (Eds A. Exelby, K. Wormington & A. Melzer) Central Queensland University, Rockhampton. Specht, R. L., Batianoff, G. N. and Reeves, R. D. (2006). Vegetation structure and biodiversity along the eucalypt forest to rainforest continuum on the serpentinite soil catena in a subhumid area of Central Queensland, Australia. Austral Ecology 31: 394-407. Specht, R. L., Salt, R. B. and Reynolds, Sally (1977). Vegetation in the vicinity of Weipa, North Queensland. Proceedings of the Royal Society of Queensland 88: 17-38. Specht, R.L. (1987). Environmental planning and management for mining and energy. Pp. 1-10. In: North Australian Mine Rehabilitation Workshop. No. 10 (Darwin, June 1986). Environmental Planning for Mining and Energy. (Eds P. J. R. Broese van Groenou and J. R. Burton). Northern Territory Dept Mines and Energy, Darwin.

6.3. Soft-rot in electricity poles after the 1974 Brisbane flood After the 1974 Brisbane Flood, the fast rate of decay of the bases of the electricity poles in the area became a concern for the Queensland Electricity Commission. On the advice of Prof. John Levy of Imperial College, London, a small Wood Poles committee was formed under Keith Cokley of the

49 Department of Forestry, Ray Staples of the Queensland Electricity Commission and Ray Specht and Mat Darveneza of the University of Queensland.

Liam Leightley, from Portsmouth, U. K., was appointed Post-doctoral Fellow in the Botany Department, University of Queensland to study this problem. A long-term experiment was established in a seasonally waterlogged site to study the rate of soft-rot in poles of local eucalypt species of north- eastern New South Wales — a forestry region that had supplied a million or more transmission poles to the Brisbane area (an region that is lacking in suitable native trees).

6.4. Growing trees on the Brisbane Tuff Early in the 1930s, thanks to the Mayne Bequest, the University of Queensland acquired the St Lucia Campus. The Head of the Botany Department, Dr Desmond H. Herbert, and Mr Ernest W. Bick, Director of the Brisbane Botanic Gardens, established a nursery of trees to be planted at appropriate positions on the campus-plan. Poinciana (Delonix regia) and other colourful deciduous trees were planted in front of the Main Building (now the Forgen-Smith Building) that was to be built on the highest level of the campus. Unfortunately, a very hard rock, the Brisbane Tuff, occurred on this site, the soil was shallow and the tree roots could not penetrate down any cracks. Many trees died, while others became very stunted, one of which has persisted into the 21st Century.

After the end of the Pacific War in 1945, the American Army volunteered to flatten the Great Court behind The Main Building.

Seedlings from a single capsule of a hybrid eucalypt, that had been studied by Dr H. Trevor Clifford, were planted along the Central Avenue of the Great Court; this resulted in a heterogeneous collection of trees. Deciduous trees flourished on the deep soil on the eastern side of the Great Court, but not on the exposed Brisbane Tuff on the western side. Buildings and Grounds dug square pits in the tuff so that trees could be planted in deep soil. These trees flourished until torrential summer storms waterlogged the holes, spectacularly during the floods of January 1974. The Botany Department then advised Buildings and Grounds to plant native eucalypts — that still flourish today.

This hard Brisbane Tuff outcrops from near the Prince Charles Hospital south to the Kangaroo Point Cliffs and west towards the St Lucia Campus. In 2006, Ray Specht visited the Lands Department archives to scan the 1840 Land Surveys of the whole Brisbane area to assess the density of eucalypt trees on the Brisbane Tuffs and on the widespread Bunya Phyllites. He was surprised to learn that a few small trees had managed to establish on the Brisbane Tuff — with densities much lower than on the Bunya Phyllites (Specht and Specht 2007). It is thus clear that shallow-rooted garden-plants would easily grow on the Brisbane Tuff, whereas deep-rooted trees and shrubs may not find cracks for their roots.

50 Reference Specht, R. L. and Specht, Alison (2007). Pre-settlement tree density in the eucalypt open-forest on the Brisbane Tuff. Proceedings of the Royal Society of Queensland 113: 9-16.

6.5. Mt Coot-tha Botanic Gardens In 1969, Brisbane Lord Mayor Clem Jones wished to establish a second Botanic Gardens to further the education of the general public. As this proposal was not supported by the Councillors who declared that the Brisbane City Botanic Gardens was rarely used, Clem opened its gates to allow motorists to drive through the Gardens on the week-ends. This was a very popular strategy that convinced the Councillors to support his motion.

The suggestion to locate the second Botanic Gardens on the Bunya Phyllites below Mt Coot-tha upset the keen rose and camellia societies who wished to locate the new Botanic Gardens on the rich soils at Long Pocket — a site frequently submerged during Brisbane River floods.

The botanists, Selwyn Everist, Len Webb and Ray Specht, who had been appointed to Clem’s Mt Coot- tha Gardens Planning Committee, stressed that most of the home gardens in the Greater Brisbane Area were on Bunya Phyllites. The new Gardens must demonstrate many different ways in which home gardens could be developed — Cactus gardens, scent gardens, rose gardens, ferneries, Japanese gardens, Camellia gardens, native flora gardens, ‘fossil’ gardens, water gardens, etc. Orchard plots of various fruit trees and vegetable gardens would introduce children to the source of these foods. Patches of lawns of different species — that could be walked across — were a valuable teaching experience.

The central attraction of the second Botanic Gardens was the spectacular geodesic-dome glasshouse housing the waterlily Victoria amazonica syn. V. regia with leaves large enough for toddlers to walk on. The waterlily had initially been collected on the Amazon River by the German naturalist, Robert Hermann Schomburgk, brother of the Director of the Adelaide Botanic Gardens. This waterlily had been a prize plant in the Brisbane Botanic Gardens, established 1855, but was washed away during the 1893 floods!

51 Inscription on a commemorative cairn in the Mt Coot-tha Botanic Gardens

BRISBANE CITY COUNCIL This cairn commemorates the commencement of the project of a new BOTANIC GARDENS to be developed in this park. 31-3-70 Clem Jones, Lord Mayor

NEW BOTANIC GARDENS COMMITTEE

Alderman C. J. GREENFIELD Mrs U. PRENTICE Alderman J. J. CAREY Alderman L. S. RUDD Mr. J. C. SLAUGHTER Mr. S. B. WATKINS Mr. I. C. HAWES Mrs S. S. LANGMORE Mr. H. W. CAULFIELD Mr. R. P. WADE Mr. G. W. BARLOW Mr. A. I. F. MacKILLOP Mr. Justice C. G. WANSTALL Mrs BELLE TAYLOR Dr. L. J. WEBB Mrs E. NALL Professor R. L. SPECHT Mrs B. R. van der BROEK Mr. S. L. EVERIST Mr. R. J. STEWARD

7. CONCLUSIONS

Consequences of global warming A lifetime of study has enabled Ray Specht to make some generalisations about the consequences of global warming for Australian ecosystems in particular. Experimental studies have shown that an increase in Global Warming of 2oC will increase aerodynamic fluxes in the atmosphere as air flows over a plant community, causing a reduction of Foliage Projective Cover and the fixation of solar energy by the foliage canopy and, thus, the species richness of the component plants and associated vertebrates (and probably invertebrates in the litter layer) in an ecosystem. In any region, there is a tendency for global warming to be associated with a change in plant community structure along the following gradient:

Open-forests —> woodlands —> tall shrublands —> low shrublands

52 About 10 million years ago, a period of Global Warming of 2-5oC caused aridity to extend from central coastal Western Australia across to south-west New South Wales. At the same time, higher rainfall in the north leached lateritic podsols to form lateritic earths (often containing bauxite), while, in the Centre, the silica in the leaf litter of Spinifex (Triodia spp.) was leached to form silcretes.

These changes in rainfall and humidity would have affected the vertical growth of foliage shoots, as well as Foliage Projective Cover.

Much more research is needed so that the countless number of producers, consumers and decomposers in each ecosystem may survive into perpetuity: —

(1) Establishment of conservation reserves for all 397 Australian ecosystems identified by TWINSPAN analysis – essential to protect for posterity the majority of our vascular plants (producers) and associated vertebrates (consumers) and invertebrates (consumers and decomposers). (2) Confirmation that the physico-chemical processes that enable solar energy, fixed by the Foliage Projective Cover, control the Alpha Biodiversity of plants and associated vertebrates in each ecosystem. (3) Understanding of why the evolution of new species may be rare in the major Australian ecosystems that appear to have retained their Alpha Biodiversity intact for millions of years. (4) Investigation of the proposition that, ten million years ago, droughts in south-western Australia confined heathy and mallee vegetation to well-watered ‘oases’ where speciation occurred, thus increasing Gamma Biodiversity in the region when the climate improved. (5) Investigation of the proposition that, during the last million years, two severe Ice Ages had lowered the sea-level, exposing the Great Barrier Reef in north Queensland, while at the same times, drought confined the rainforest to ‘oases’ where speciation occurred, thus increasing Gamma Biodiversity in the region.

8. Publications of Raymond Louis Specht

(1) Books

1. Specht, R. L. and Mountford, C. P. (Eds) (1958). Records of the American-Australian Expedition to Arnhem Land. Vol. 3. Botany and Plant Ecology. Melbourne University Press, Melbourne, 522 pp. 2. Specht, R. L. (Ed.) (1964). Records of the American-Australian Expedition to Arnhem Land. Vol. 4. Zoology. Melbourne University Press, Melbourne, 533 pp. Specht, R. L. and Specht, Alison (Eds) (1998). Personal Experiences on the 'Coast of Adventure'. American-Australian Scientific Expedition to Arnhem Land, March-November 1948. Arnhem Land Reunion, Sydney, 27-28th June 1998. Southern Cross University, Lismore, N.S.W. 3. Specht, R. L. (1972). The Vegetation of South Australia. Govt Printer, Adelaide, 328 pp. 4. Specht, R. L., Roe, Ethel M. and Boughton, Valerie H. (Eds) (1974). Conservation of Major Plant Communities in Australia and Papua New Guinea. Australian Journal of Botany Suppl. No. 7, 667 pp. 5. Clifford, H. T. and Specht, R. L. (1979). The Vegetation of North Stradbroke Island. (with notes on the fauna of mangrove and marine meadow ecosystems by Marion M. Specht). University of Queensland Press, St Lucia, 141 pp. 6. Specht, R. L. (Ed.) (1979). Ecosystems of the World. Vol. 9A. Heathlands and Related Shrublands. Descriptive Studies. Elsevier, Amsterdam, 497 pp.

53 7. Specht, R. L. (Ed.) (1981). Ecosystems of the World. Vol. 9B. Heathlands and Related Shrublands. Analytical Studies. Elsevier, Amsterdam, 385 pp. 8. Di Castri, F., Goodall, D. W. and Specht, R. L. (Eds) (1981). Ecosystems of the World. Vol. 11. Mediterranean-type Shrublands. Elsevier, Amsterdam, 650 pp. 9. Specht, R. L. (Coord.) (1981). Part 2. Flora. In: Ecological Biogeography of Australia.Vol. 1. 805 pp. (Ed. A. Keast). Junk, the Haag 10. Clifford, H.T. and Specht, R.L. (Eds) (1986). Tropical Plant Communities. Their Resilience, Function and Management in Northern Australia. Utah Foundation & Bot. Dept, Univ. Qld, St Lucia, Qld, 217pp. 11. Specht, R.L. (Ed.) (1988). Mediterranean-type Ecosystems. A Data Source Book. (Tasks for Vegetation Sciences. Vol. 19). Kluwer Academic Publ., Dordrecht, 248 pp. 12. Specht, R. L., Specht, Alison, Whelan, M. B. and Hegarty, Elwyn E. (1995). Conservation Atlas of Plant Communities in Australia. Centre for Coastal Management, Southern Cross University, Lismore, N.S.W., 1073 pp. 13. Specht, R. L. and Specht, Alison (1999 Hardback; 2002 Paperback). Australian Plant Communities. Dynamics of Structure, Growth and Biodiversity. Oxford University Press, Melbourne, 492 pp. 14. Specht, R. L., Wahlqvist, M. L., Wahlquist, G. & Specht, G. J. (Eds) (2009). The Flight of the Woodpecker. The migration of five families from Germany to South Australia. Published by Gil Wahlquist in Microsoft Windows. 15. Thomas, M. & Neale, Margo (Eds). (2011). Exploring the Legacy of the 1948 Arnhem Land Expedition. Australian National University E Press, Canberra.

(2) Australian plant communities — Ecological studies

16. Specht, R. L. and Perry, R. A. (1948). Plant ecology of part of the Mount Lofty Ranges. Transactions of the Royal Society of South Australia 72: 91-132. 17. Specht, R. L. (1951). A reconnaissance survey of the soils and vegetation of the Hundreds of Tatiara, Wirrega and Stirling. Transactions of the Royal Society of South Australia 74: 79-107. 18. Specht, R. L. (1958). Gymnospermae and angiospermae collected on the Arnhem Land Expedition. (including 20 new taxa) Pp. 185-318. In: Records of the American-Australian Expedition to Arnhem Land. Vol. 3. Botany and Plant Ecology. (Eds R. L. Specht and C. P. Mountford). Melbourne University Press, Melbourne. 19. Specht, R. L. (1958). Climate, geology, soils and plant ecology of the northern portion of Arnhem Land. Pp. 333- 414. In: Records of the American-Australian Expedition to Arnhem Land. Vol. 3. Botany and Plant Ecology. (Eds R. L. Specht and C. P. Mountford). Melbourne University Press, Melbourne. 20. Specht, R. L. and W. Bateman (1958). Some ecological and systematic notes on the vegetation near Mt Marumba in the centre of Arnhem Land. Pp. 327-331. In: Records of the American-Australian Expedition to Arnhem Land. Vol. 3. Botany and Plant Ecology. (Eds R. L. Specht and C. P. Mountford). Melbourne University Press, Melbourne. 21. Specht, R. L., Brownell, P. F. and Hewitt, P. N. (1961). The plant ecology of the Mount Lofty Ranges, South Australia. 2. The distribution of Eucalyptus elaeophora. Transactions of the Royal Society of South Australia 85: 155-176. 22. Specht, R. L. (1962). The nomenclature of some Borreria species from Northern Australia. Kew Bulletin 16: 239-240. 23. Carrodus, B. B., Specht, R. L. and Jackman, Margaret E. (1965). The vegetation of Koonamore Station, South Australia. Transactions of the Royal Society of South Australia 89: 41-57. 24. Specht, R. L. (1969). The vegetation of Pearson Islands. A re-examination - February 1960. Transactions of the Royal Society of South Australia 93: 143-152. 25. Specht, R. L. (1970). Vegetation. Pp. 44-67. In The Australian Environment. 4th Edition. (Ed. G. W. Leeper) C.S.I.R.O. - Melbourne University Press, Melbourne. 26. Specht, R. L., Roe, Ethel M. and Boughton, Valerie. H. (Eds) (1974). Conservation of major plant communities in Australia and Papua New Guinea. Australian Journal of Botany Supplement No. 7, 667 pp. 27. Specht, R. L. (1975). Stradbroke Island: A place for teaching biology. Proceedings of the Royal Society of Queensland 86: 81-83. 28. Shackleton, N. J. & Kennett, J. P. (1975). Palaeo-temperature history of the Cenozoic and the initiation of Antarctic glaciation: Oxygen and carbon isotope analyses in DSDP sites 277, 279, 281. Pp. 743-755. In: Initial Reports of the Deep Sea Drilling Project, 29. United States Government Printing Office, Washington. 29. Specht, R. L., Salt, R. B. and Reynolds, Sally (1977). Vegetation in the vicinity of Weipa, North Queensland. Proceedings of the Royal Society of Queensland 88: 17-38. 30. Byrnes N., Everist, S. L., Reynolds, Sally, Specht, Alison and Specht, R. L. (1977). Vegetation of Lizard Island, North Queensland. Proceedings of the Royal Society of Queensland 88: 1-15. 31. Clifford, H. T. and Specht, R. L. (1979). The Vegetation of North Stradbroke Island. (with notes on the fauna of mangrove and marine meadow ecosystems by Marion M. Specht). University of Queensland Press, St Lucia, 41 pp. 32. Specht, R. L. (1979). Heathlands and related shrublands of the world. Pp 1-18. In Ecosystems of the World. Vol. 9A. Heathlands and Related Shrublands. Descriptive Studies. (Ed. R. L. Specht) Elsevier, Amsterdam. 33. Specht, R. L. (1979). The sclerophyllous (heath) vegetation of Australia: The eastern and central States. Pp. 125-210. In Ecosystems of the World. Vol. 9A. Heathlands and Related Shrublands. Descriptive Studies. (Ed. R. L. Specht) Elsevier, Amsterdam. 34. Specht, R. L. (1979). 'Maquis' vegetation of New Caledonia. Pp. 317-319. In: Ecosystems of the World. Vol. 9A. Heathlands and Related Shrublands. Descriptive Studies. (Ed. R. L. Specht) Elsevier, Amsterdam.

54 35. Specht, R. L. (1979). Alpine scrub and heathland communities in Japan. Pp. 425-426. In: Ecosystems of the World. Vol. 9A. Heathlands and Related Shrublands. Descriptive Studies. (Ed. R. L. Specht) Elsevier, Amsterdam. 36. Specht, R. L. and Womersley, J. S. (1979). Heathlands and related shrublands of Malesia (with particular reference to Borneo and New Guinea). Pp. 321-338. In: Ecosystems of the World. Vol. 9A. Heathlands and Related Shrublands. Descriptive Studies. (Ed. R. L. Specht) Elsevier, Amsterdam. 37. Specht, R. L. (1981). Major vegetation formations in Australia. Pp. 163-298. In: Ecological Biogeography of Australia. (Ed. A. K east). Junk, the Haag. 38. Specht, R. L. (1981). Mallee ecosystems in southern Australia. Pp. 203-231. In Ecosystems of the World. Vol. 11. Mediterranean-type Shrublands. (Eds F. di Castri, D. W. Goodall and R. L. Specht). Elsevier, Amsterdam. 39. Specht, R. L. (1981). Heathlands. Pp. 253-275. In: Australian Vegetation. (Ed. R. H. Groves). Cambridge University Press, Cambridge. 40. Specht, R. L. (1981). Structural attributes - foliage projective cover and standing biomass. Pp. 10-21. In Vegetation Classification in the Australian Region. (Eds A. N. Gillison and D. J. Anderson). C.S.I.R.O. and Australian National University Press, Canberra. 41. Groves, R. H. and Specht, R. L. (1981). Seral considerations in heathland. Pp. 78-85. In: Vegetation Classification in the Australian Region. (Eds A. N. Gillison and D. J. Anderson). C.S.I.R.O. and Australian National University Press, Canberra. 42. Specht, R. L. (July 1981). Heathlands: An introduction. pp. 7-12. In Heaths in New South Wales. Parks and Wildlife New South Wales, Sydney. 43. Specht, R. L. (1982). General characteristics of Mediterranean-type ecosystems. Pp. 13-19. In : Proceedings of the Symposium on Dynamics and Management of Mediterranean-type Ecosystems, June 22-26, 1981, San Diego, California. (Eds C. E. Conrad and W. C. Oechel). U.S.D.A. Forest Service, General Technical Report PSW-58, Washington, D.C. 44. Specht, R. L. and Moll, E. J. (1983). Mediterranean-type heathlands and sclerophyllous shrublands of the world - An overview. Pp. 41-65. In: Mediterranean-Type Ecosystems. The Rôle of Nutrients. (Eds F. J. Kruger, D. T. Mitchell and J. U. M. Jarvis). Springer-Verlag, Berlin. 45. Groves, R. H., Beard, J. S., Deacon, H. J., Lambrechts, J. J. N., Rabinovitch-Vin, A., Specht, R. L. and Stock, W. D. (1983). Introduction: The origins and characteristics of mediterranean ecosystems. Pp. 1-17. In: Mineral Nutrients in Mediterranean Ecosystems. (Ed. J. A. Day). South African National Scientific Programmes Report No. 71. 46. Specht, R.L. (1986). Tropical and subtropical plant communities in Australia. Pp. 3-11. In: Tropical Plant Communities. (Eds H.T. Clifford and R.L. Specht). Bot. Dept., Univ. Qld, St Lucia, Qld. 47. Specht, R.L. (1986). Heathlands-heathy shrublands: Dry heathland on deep Quaternary sand. P. 145. In: Tropical Plant Communities. (Eds H.T. Clifford and R.L. Specht). Bot. Dept., Univ. Qld, St Lucia, Qld. 48. Specht, R.L. (1986). Forested wetland vegetation: Melaleuca leucadendron/M. quinquenervia. p. 151. In: Tropical Plant Communities. (Eds H.T. Clifford and R.L. Specht). Bot. Dept., Univ. Qld, St Lucia, Qld. 49. Specht, R.L. (1986). Coastal dune vegetation: Low closed-forest or coastal woodland. P. 153. In: Tropical Plant Communities. (Eds H.T. Clifford and R.L. Specht). Bot. Dept., Univ. Qld, St Lucia, Qld. 50. Catling, P.C., Daget, Ph., Fox, B.J., Greenslade, Penny, Majer, J.D., Orshan, G., Rundel, P.W., Specht, R.L. and Westman, W.E. (1988). Climate, vegetation, vertebrates and soil/litter invertebrates of mediterranean-type ecosystems - Data banks. Pp. 3-9. In: Mediterranean-type Ecosystems. A Data Source Book. (Ed. R.L. Specht). Kluwer Academic Publ., Dordrecht. 51. Specht, R.L. (1990). Forested wetlands in Australia. Pp.387-406. In: Ecosystems of the World. Vol. 16. Wetland Forests. (Eds A.E. Lugo, M.M. Brinson and Sandra Brown). Elsevier, Amsterdam. 52. Specht, R.L, Clifford, H.T., Arianoutsou, Margarita, Bird, L.H., Bolton, M.P., Forster, P.I., Grundy, Rhonda.I., Hegarty, Elwyn E. and Specht, Alison (1991). Structure, floristics and species richness of plant communities in southeast Queensland. Proceedings of the Royal Society of Queensland 101 : 27-78. 53. Batianoff, G.N., Specht, R.L. and Reeves, R.D. (1991). The serpentinite flora of the humid subtropics of eastern Australia. Proceedings of the Royal Society of Queensland 101 : 137-158. 54. Specht, R.L. (1993). Vegetation. Pp. 91-117. In: Land Degradation Processes in Australia. (Eds G. H. McTainsh and W. C. Boughton). Longman Cheshire, Melbourne. 55. Specht, R.L. (1994). Dry coastal ecosystems of Australia. An overview of the dune vegetation. Pp. 223 - 237. In: Ecosystems of the World. Vol. 2 B. Dry Coastal Ecosystems: Africa, America, Asia and Oceania. (Ed. E. van der Maarel). Elsevier, Amsterdam. 56. Specht, R. L. (1994). Heathlands. Pp. 321-344. In: Australian Vegetation. Second Edition. (Ed. R. H. Groves). Cambridge University Press, Cambridge. 57. Specht, Alison and Specht, R. L. (2000). Australia: Biodiversity of ecosystems. pp. 307-324. In: Encyclopedia of Biodiversity. (Ed. S. Levin) Academic Press, San Diego, California. 58. Specht, R. L. and Specht, Alison (2002). Objective classification of plant communities in tropical and subtropical Australia. Proceedings of the Royal Society of Queensland 110: 65-82. 59. Martin, Helene A. and Specht, R. L. (2005). Sclerophyll (heathy) understoreys in the Mount Lofty Ranges, South Australia. Transactions of the Royal Society of South Australia 129: 14-24. 60. Specht, R. L., Batianoff, G. N. and Reeves, R. D. (2005). The vegetation continuum on the serpentinite soil catena in the Rockhampton – Marlborough area. Pp. 23-40. In: Marlborough Serpentine, Landscape Management and Conservation. (Eds Anne Exelby, K. Wormington and A. Melzer) Centre for Environmental Management, Central Queensland University, Rockhampton. 61. Specht, R. L. (2012). Plant communities of North Stradbroke Island: Development of structure and species richness. Proceedings of the Royal Society of Queensland 117: 187-197.

55

(3) Australian plant communities — Conservation 62. Specht, R. L. (1953, reprinted 1964). The vegetation of the National Park and Reserves. Pp. 29-58 In National Park and Reserves. Government Printer, Adelaide. 63. Specht, R. L. and Cleland, J. B. (1961). Flora conservation in South Australia. 1. The preservation of plant formations and associations recorded in South Australia. Transactions of the Royal Society of South Australia 85: 177-196. 64. Specht, R. L. and Cleland, J. B. (1964). Flora conservation in South Australia. 2. The preservation of species recorded in South Australia. Transactions of the Royal Society of South Australia. 87: 63-92. 65. Specht, R. L. (May 1971). I.B.P. studies on productivity and conservation of terrestrial ecosystems. Australian Science Teachers' Journal 17(1): 33-36. 66. Specht, R. L., Roe, E. M. and Boughton, V. H. (Eds) (1974). Conservation of major plant communities in Australia and Papua New Guinea. Australian Journal of Botany Suppl. No. 7, 667 pp. 67. Specht, R. L. (1975). A national system of ecological reserves in Australia. The report and its recommendations. Pp. 11-21 In: A National System of Ecological Reserves in Australia. (Ed. F. Fenner) Australian Academy of Science Report No. 19. 68. Specht, R. L. (1977). Conservation by cultivation. Pp. 3-5. In: Growing Australian Native Plants. (Ed. R. Prater) Dept. of Continuing Education (Namoi), University of New England, Tamworth, N.S.W. 69. Specht, R. L. (1978). Conditions for cultivation of Australian plants. Australian Plants 9: 312-313. 70. Specht, R. L. (July 1978). In Wildness is the Preservation of the World. Sixth Romeo Watkins Lahey Memorial Lecture, National Parks Association of Queensland, Brisbane, 12pp. 71. Specht, R. L. (1981). Conservation: Australian heathlands. Pp. 235-240. In: Ecosystems of the World. Vol. 9B. Heathlands and Related Shrublands. Analytical Studies. (Ed. R. L. Specht). Elsevier, Amsterdam. 72. Specht, R. L. (1981). Conservation of vegetation types. Pp. 393-410. In: Australian Vegetation. (Ed. R. H. Groves). Cambridge University Press, Cambridge. 73. Allen, R., Bowmaker, A. P., Brain, C. K., Edwards, D., Newman, G., Owen-Smith, R. N., Ribbink, A. J. and Specht, R. L. (1982). Goals of ecosystem conservation. Pp. 4-9. In: Conservation of Ecosystems: Theory and Practice. (Eds W. R. Siegfried and B. R. Davies). South African National Scientific Programmes Report No. 61. 74. Bodkin, D. B., Davies B. R., Edwards, D., Frost, P. G. H., Lucas, G. L., Newman, G., Rabinovitch-Vin, A., Ray, G. C., Specht, R. L., van der Elst, R. and Walker, B. H. (1982). Ecological characteristics of ecosystems. Pp. 17-27: In: Conservation of Ecosystems: Theory and Practice. (Eds W. R. Siegfried and B. R. Davies). South African National Scientific Program Report No. 61. 75. Bolton, M. P. and Specht, R. L. (1983). A Method for Selecting Nature Conservation Reserves. Australian National Parks and Wildlife Service (Canberra) Occasional Paper No. 8, 32 pp. 76. Purdie, R. W. (1987). Selection of key area networks for regional nature conservation — the revised Bolton and Specht Method. . Proceedings of the Royal Society of Queensland 97: 59-71. (World Wildlife (Aust.) Fund grants to R. L. Specht and Queensland National Parks.) 77. Specht, R. L. (1985). Nutritional problems of native plants (Bill Tulloch Memorial Lecture). Society for Growing Australian Plants (Qld Region) Bulletin 24(2): 16-20. 78. Specht, R. L. (1990). Comment on conserving what? The basis for nature conservation reserves in New South Wales 1967-1988. Australian Zoologist 6(2): 76-77. 79. Specht, Alison and Specht, R. L. (1993). Species richness and canopy productivity of Australian plant communities. Biodiversity and Conservation 2: 152 - 167. 80. Specht, Alison and Specht, R. L. (1994). Biodiversity of overstorey trees in relation to canopy productivity and stand density in the climatic gradient from warm temperate to tropical Australia. Biodiversity Letters 2: 39- 45. 81. Specht, R. L. (1994). Biodiversity and conservation. Pp. 525-555. In: Australian Vegetation. Second Edition. (Ed. R. H. Groves). Cambridge University Press, Cambridge. 82. Specht, R. L. and Specht, Alison (October 1994). Conservation of plant biodiversity in the Mediterranean-type climate of southern Australia. MEDECOS Resúmenes. Noticiero de Biologia, Chile 2(3): 53. 83. Specht, R. L., Specht, Alison, Whelan, M. B. and Hegarty, Elwyn E. (1995). Conservation Atlas of Plant Communities in Australia. Centre for Coastal Management, Southern Cross University, Lismore, N.S.W., 1073 pp. 84. Specht, R. L. and Specht, Alison (2000). Australia: Biodiversity of ecosystems. Pp. 307-324. In: Encyclopedia of Biodiversity. (Ed. S. Levin) Academic Press, San Diego, California. 85. Stubbs, B. J. and Specht, R. L. (2005). Lamington National Park and Binna Burra Mountain Lodge, Queensland: Partners in conserving rainforest. Pp. 707-716. In: Proceedings 6th National Conference of the Australian Forest History Society. (Eds M. Calver et al.) Millpress, Rotterdam.

(4) Australian plant communities — Ecological Biogeography

86. Specht, R. L. (1958). Geographical relationships of the flora of Arnhem Land. Pp. 415-478. In: Records of the American-Australian Expedition to Arnhem Land. Vol. 3. Botany and Plant Ecology. (Eds R. L. Specht and C. P. Mountford). Melbourne University Press, Melbourne. 87. Specht, R. L., (Co-ordinator) (1981). The flora of Australia. Pp. 163-805. In: Ecological Biogeography of Australia...Vol. 1 (Ed. A. Keast). Junk, the Haag. 88. Specht, R. L. (1981). Biogeography of halophytic angiosperms (salt-marsh, mangrove, sea-grass). Pp. 575-590. In: Ecological Biogeography of Australia. (Eds A. Keast et al.). Junk, the Haag.

56 89. Specht, R. L. (1981). Evolution of the Australian flora: Some generalizations. Pp. 783-806. In : Ecological Biogeography of Australia. (Eds A. Keast et al.). Junk, the Haag. 90. Specht, R.L. (1988). Origin and evolution of terrestrial plant communities in the wet-dry tropics of Australia. Proceedings of the Ecological Society of Australia 15: 19-30. 91. Specht, R.L. , Dettmann, Mary E. and Jarzen, D. M. (1992). Community associations and structure in the Late Cretaceous vegetation of southeast Australasia and Antarctica. Palaeogeography, Palaeoclimatology, Palaeoecology 94 : 283-309. 92. Specht, R.L. and Dettmann, Mary.E. (1995). Palaeo-ecology of Australia and current physiological functioning of plant communities. Pp. 201-214. In: Time Scales of Biological Responses to Water Constraints: The Case of Mediterranean Biota. (Eds J. Roy, J. Aronson and F. di Castri) SPB Academic Publishing, Amsterdam. 93. Specht, A. and Specht, R. L. (2005). Historical biogeography of Australian forests. Pp. 1-7. In: Australia and New Zealand Forest Histories. (Ed. J. Dargavel) Australian Forest History Society Inc. Occasional Publication No. 1.

(5) Australian plant communities — Community physiology

94. Specht, R. L. and Rayson, Patricia (1957). Dark Island heath (Ninety-Mile Plain, South Australia). 1. Definition of the ecosystem. Australian Journal of Botany 5: 52-85. 95. Rayson, Patricia (1957). Dark Island heath (Ninety-Mile Plain, South Australia). 2. The effects of micro- topography on climates, soils and vegetation. Australian Journal of Botany 5: 86-102.(M. Sc. postgraduate studies, supervised by R. L. Specht) 96. Specht, R. L. and Rayson, Patricia (1957). Dark Island heath (Ninety-Mile Plain, South Australia). 3. The root systems. Australian Journal of Botany 5: 103-114. 97. Specht, R. L. (1957). Dark Island heath (Ninety-Mile Plain, South Australia). 4. Soil moisture patterns produced by rainfall interception and stem-flow. Australian Journal of Botany 5: 137-150. 98. Specht, R. L. (1957). Dark Island heath (Ninety-Mile Plain, South Australia). 5. The water relationships in heath vegetation and pastures on the Makin Sand. Australian Journal of Botany 5: 151-172. 99. Specht, R. L., Rayson, Patricia and Jackman, Margaret E. (1958). Dark Island heath (Ninety-Mile Plain, South Australia). 6. Pyric succession: Changes in composition, coverage, dry weight, and mineral nutrient status. Australian Journal of Botany 6: 59-88. 100. Specht, R. L. (1958). Micro-environment (soil) of a natural plant community. Arid Zone Research (UNESCO) 11: 152-155. 101. Specht, R. L. (1958). Range management in the arid zone of the United States. Arid Zone Newsletter 1958: 75- 78. 102. Martin, Helene A. and Specht, R. L. (1962). Are mesic communities less drought resistant? A study on moisture relationships in dry sclerophyll forest at Inglewood, South Australia. Australian Journal of Botany 10: 106- 118. 103. Specht, R. L. (1963). Dark Island heath (Ninety-Mile Plain, South Australia). 7. The effect of fertilizers on composition and growth, 1950-60. Australian Journal of Botany 11: 67-94. 104. Hall, E. Anne A., Specht, R. L. and Eardley, Constance M. (1964). Regeneration of the vegetation on Koonamore Vegetation Reserve, 1926-62. Australian Journal of Botany 12: 205-264. 105. Groves, R. H. and Specht, R. L. (1965). Growth of heath vegetation. 1. Annual growth curves of two heath ecosystems in Australia. Australian Journal of Botany 13: 261-280. 106. Carrodus, B. B. and Specht, R. L. (1965). Factors affecting the relative distribution of Atriplex vesicaria and Kochia sedifolia (Chenopodiaceae) in the arid zone of South Australia. Australian Journal of Botany 13: 419-433. 107. Specht, R. L. and Groves, R. H. (1966). A comparison of the phosphorus nutrition of Australian heath plants and introduced economic plants. Australian Journal of Botany 14: 201-221. 108. Brownell, P. F. (1965). Sodium as an essential micro-nutrient element for a higher plant (Atriplex vesicaria). Plant Physiology 40: 460-468. 109. Jeffrey, D. W. (1964). The formation of polyphosphate in Banksia ornata, an Australian heath plant. Australian Journal of Biological Science 17: 845-854. 110. Jeffrey, D. W. (1967). Phosphate nutrition of Australian heath plants. 1. The importance of proteoid roots in Banksia (Proteaceae). Australian Journal of Botany 15: 403-412 111. Jeffrey, D. W. (1968). Phosphate nutrition of Australian heath plants. II. The formation of polyphosphate by five heath species. Australian Journal of Botany 16: 603-613. 112. Haines, Wendy M. (1967). Experimental Approaches to the Study of Seasonal Root Activity and Phosphorus Nutrition of Australian Heath Plants. M. Sc. Thesis, University of Melbourne. 113. Specht, R. L. (1966). The growth and distribution of mallee-broombush (Eucalyptus incrassata - Melaleuca uncinata association) and heath vegetation near Dark Island Soak, Ninety-Mile Plain, South Australia. Australian Journal of Botany 14: 361-371. 114. Specht, R. L. (1966). The International Biological Program (I. B. P.) Section PCT: Productivity, production processes and conservation of terrestrial communities. Australian Journal of Science 28: 377-380. 115. Specht, R. L. (1967). The International Biological Programme. Aust. Science Teachers' Journal 13(3): 43-48. 116. Jones, R. and Specht, R. L. (1967). Productivity studies on heath vegetation in southern Australia. A comment on sampling problems. Folia Geobot. Phytotax. Praha 2: 337-346. 117. Parsons, R. F. and Specht, R. L. (1967). Lime chlorosis and other factors affecting the distribution of Eucalyptus on coastal sands in southern Australia. Australian Journal of Botany 15: 95-105.

57 118. Specht, R. L. (1967). The photosynthesis of plant communities in relation to structure, physiology and environment. Photosynthetica 1: 132-134. 119. Specht, R. L. and Angus, D. E. (1967). The effect of community structure and physiology in the transformation of solar energy. Journal of the. Australian Institute of Agricultural Science 33: 229-230. 120. Specht, R. L. (1969). A comparison of the sclerophyllous vegetation characteristic of Mediterranean-type climates in France, California and southern Australia. 1. Structure, morphology and succession. Australian Journal of Botany 17: 277-292. 121. Specht, R. L. (1969). A comparison of the sclerophyllous vegetation characteristic of Mediterranean-type climates in France, California and southern Australia. 2. Dry matter, energy and nutrient accumulation. Australian Journal of Botany 17: 293-308. 122. Jones, R., Groves, R. H. and Specht, R. L. (1969). Growth of heath vegetation. 3. Growth curves for heath in southern Australia. A reassessment. Australian Journal of Botany 17: 309-314. 123. Specht, R. L. and Jones, R. (1971). A comparison of the water use by heath vegetation at Frankston, Victoria and Dark Island Soak, South Australia. Australian Journal of Botany 19: 311-326. 124. Specht, R. L. (1972). Water use by plant communities in the arid zone of Australia. Pp. 48-53. In ::Eco- physiological Foundation of Ecosystem Productivity in the Arid Zone. Publishing House "Nauka", Leningrad. 125. Specht, R. L. (1972). Water use by perennial, evergreen plant communities in Australia and Papua New Guinea. Australian Journal of Botany 20: 273-299. 126. Specht, R. L. (1972). The Vegetation of South Australia. Govt Printer, Adelaide, 328 pp. 127. Specht, R. L. (1973). Structure and functional response of ecosystems in the mediterranean climate of Australia. Pp. 113-120. In: Mediterranean Type Ecosystems. Origin and Structure. (Eds. F. di Castri and H. A. Mooney) Springer-Verlag, Berlin. 128. Specht, R. L. and Brouwer, Yvonne M. (1975). Seasonal shoot growth of Eucalyptus spp. in the Brisbane area of Queensland (with notes on shoot growth and litter fall in other areas of Australia). Australian Journal of Botany 23: 459-474. 129. Specht, R. L., Brouwer, Yvonne M. and Derrick, E. H. (1975). Seasonal waves of asthma: A possible botanical cause. International Journal of Biometeorology 19: 28-36. 130. Specht, R. L. (1975). A heritage inverted, our flora endangered. Search 6: 472-477. 131. Heddle, Elizabeth M. and Specht, R. L. (1975). Dark Island heath (Ninety-Mile Plain, South Australia). 8. The effect of fertilizers on composition and growth, 1950-1972. Australian Journal of Botany 23: 151-164. 132. Specht, R. L. (1975). The effect of fertilizers on sclerophyll (heath) vegetation. The problems of revegetation after sand-mining of high dunes. Search 6: 459-461. 133. Specht, R. L. (1976). Heath soils and phosphorus. Search 7: 133. 134. Specht, R. L. (1977). The effects of fire on the Fraser Island ecosystems. University of Queensland Anthropology Museum, Occasional Papers in Anthropology No. 8: 139-146. 135. Specht, R. L. (1976). Ranger Uranium Environmental Inquiry. Sydney, June 16th 1976. Submission prepared by R. L. Specht, with assistance from: I. L. Ophel, Atomic Energy of Canada, Chalk River, Canada Dr D. Doley, Botany Dept., University of Queensland

i. The Arnhem Land Escarpment: — A National Monument. ii. Sensitivity of the Sandstone Ecosystems to Environmental Change. iii. Sensitivity of the Black Soil Flood-plain Ecosystems to Environmental Change. iv. (The effect of background radiation on the speciation of unicellular algae,) v. Revegetation and Weeds. 132 Saenger, P., Specht, Marion M., Specht, R. L. and Chapman, V. J. (1977). Mangal and coastal salt marsh communities in Australasia. Pp. 293-345 In: Ecosystems of the World. Vol. 1. Wet Coastal Formations. (Ed. V. J. Chapman). Elsevier Scientific Publ. Co., Amsterdam. 133 Specht, R. L., Connor, D. J. and Clifford, H. T. (1977). The heath-savannah problem: The effect of fertilizer on sand-heath vegetation of North Stradbroke Island, Queensland. Australian Journal of Ecology 2: 179- 186. 134 Edmonds, S. J. and Specht, Marion M. (1981). Dark Island heathland, South Australia: Faunal rhythms Pp. 15-27. In: Ecosystems of the World. Vol. 9B. Heathlands and Related Shrublands. Analytical Studies. (Ed. R. L. Specht). Elsevier, Amsterdam. (Co-workers in the study of heathland ecosystems). 135 Specht, Marion M. (1991). Seasonality of aerial, litter and soil fauna in Dark Island heathland, South Australia. Pp. 39-42. In: Soil and Litter Invertebrates of Australian Mediterranean-Type Ecosystems. (Eds Penny Greenslade and J. D. Majer) Western Australian Institute of Technology, School of Biology Bulletin No. 12. 136 Specht, R. L. (1981). The water relations of heathlands: Seasonal water-logging. Pp. 99-106. In: Ecosystems of the World. Vol. 9B. Heathlands and Related Shrublands. Analytical Studies. (Ed. R. L. Specht). Elsevier, Amsterdam. 137 Specht, R. L. (1981). The water relations of heathlands: Morphological adaptations to drought. Pp. 123- 129: In: Ecosystems of the World. Vol. 9B. Heathlands and Related Shrublands. Analytical Studies. (Ed. R. L. Specht). Elsevier, Amsterdam. 138 Specht, R. L., Rogers, R. W. and Hopkins, A. J. M. (1981). Seasonal growth and flowering rhythms: Australian heathlands. Pp. 5-13. In: Ecosystems of the World. Vol. 9B. Heathlands and Related Shrublands. Analytical Studies. (Ed. R. L. Specht). Elsevier, Amsterdam.

58 139 Coleman, R. G. and Specht, R. L. (1981). Mineral nutrition of heathlands: The rôle of polyphosphate in the phosphorus economy of heathland species. Pp. 197-207. In: Ecosystems of the World. Vol. 9B. Heathlands and Related Shrublands. Analytical Studies. (Ed. R. L. Specht). Elsevier, Amsterdam. 140 Ozanne, P. G. and Specht, R. L. (1981). Mineral nutrition of heathlands: Phosphorus toxicity. Pp. 209-213. In: Ecosystems of the World. Vol. 9B. Heathlands and Related Shrublands. Analytical Studies. (Ed. R. L. Specht). Elsevier, Amsterdam. 141 Specht, R. L. (1981). Ecophysiological principles determining the biogeography of major vegetation formations in Australia. Pp. 299-332. In: Ecological Biogeography of Australia. (Eds A. Keast et al.). Junk, the Haag. 142 Specht, R. L. (1981). Responses to fire in heathlands and related shrublands. Pp. 395-415. In: Fire and the Australian Biota. (Eds A. M. Gill, R. H. Groves and I. R. Noble). Australian Academy of Science, Canberra. 143 Specht, R. L. (1981). Primary production in mediterranean-climate ecosystems regenerating after fire. Pp. 257-267. In: Ecosystems of the World. Vol. 11. Mediterranean-type Shrublands. (Eds F. di Castri, D. W. Goodall and R. L. Specht). Elsevier, Amsterdam. 144 Specht, R. L. (1981). Nutrient release from decomposing leaf litter of Banksia ornata, Dark Island heathland, South Australia. Australian Journal of Ecology 6: 59-63. 145 Specht, R. L. and Morgan, D. G. (1981). The balance between the foliage projective covers of overstorey and understorey strata in Australian vegetation. Australian Journal of Ecology 6: 193-202. 146 Specht, R. L. (1981). Growth indices - Their rôle in understanding the growth, structure and distribution of Australian vegetation. Oecologia, Berlin 50: 347-356. 147 Specht, R. L. (1982). Interaction of research and management of mediterranean-type ecosystems. Pp. 562- 563. In: Proceedings of the Symposium on Dynamics and Management of Mediterranean-type Ecosystems, June 22-26, 1981, San Diego, California. (Eds C. E. Conrad and W. C. Oechel). U.S.D.A. Forest Service, General Technical Report PSW-58, Washington, D.C. 148 Specht, R. L., Moll, E. J., Pressinger, F. and Sommerville, Jacqui (1983). Moisture regime and nutrient control of seasonal growth in mediterranean ecosystems. Pp. 120-132. In: Mediterranean-Type Ecosystems. The Rôle of Nutrients. (Eds F. J. Kruger, D. T. Mitchell and J. U. M. Jarvis). Springer-Verlag, Berlin. 149 Specht, R. L. (1983). Foliage projective covers of overstorey and understorey strata of mature vegetation in Australia. Australian Journal of Ecology 8: 433-439. 150 Specht, R. L., Clifford, H. T. and Rogers, R. W. (1984). Species richness in a eucalypt open-woodland on North Stradbroke Island, Queensland. The effect of overstorey and fertilizer, 1965-1984. Pp. 267-277. In: Focus on Stradbroke. New Information on North Stradbroke Island and Surrounding Areas, 1974- 1984. (Eds R. J. Coleman, Jenny Covacevich and P. Davie). Boolarong Press, Brisbane. 151 Specht, R. L. (1984). The mediterranean climate of Australia and its impact on the structure and distribution of vegetation. Bulletin de Societé de Botanique, France 131: 225-234. 152 Specht, R. L. (1984). Species-richness of plant communities in the mediterranean climate of south-eastern Australia. Pp. 143-144. In: MEDECOS IV. Proceedings of the 4th International Conference on Mediterranean Ecosystems. (Ed. B. Dell). Botany Department, University of Western Australia, Nedlands. 153 Clifford, H.T. and Specht, R.L. (Eds) (1986). Tropical Plant Communities. Their Resilience, Function and Management in Northern Australia. Utah Foundation & Botany Dept, Univ. Qld, St Lucia, Qld, 217pp. 154 Specht, R.L. (1986). Functioning of tropical plant communities: Phenology. Pp. 78-90. In: Tropical Plant Communities. (Eds H.T. Clifford and R.L. Specht). Botany Dept., Univ. Qld, St Lucia, Qld. 155 Clifford, H.T., Charles-Edwards, D.A., Doley, D. and Specht, R.L. (1986). Plant science research in tropical- subtropical Australia. Pp. 123-132. In: Tropical Plant Communities. (Eds H.T. Clifford and R.L. Specht). Botany Dept., Univ. Qld, St Lucia, Qld. 156 Specht, R.L. (1987). The effect of summer drought on vegetation structure in the mediterranean climate region of Australia. Pp. 625-639. In:: Plant Response to Stress. Functional Analysis in Mediterranean Ecosystems. (Eds J.D. Tenhunen, F.M. Catarino, O.L. Lange and W.C. Oechel). (NATO Advanced Science Institute Series, Vol. G15) Springer-Verlag, Berlin. 157 Specht, R.L. (1987). Environmental planning and management for mining and energy. Pp. 1-10. In: North Australian Mine Rehabilitation Workshop. No. 10. (Darwin, June 1986). Environmental Planning for Mining and Energy. (Eds P.J.R. Broese van Groenou and J.R. Burton). Northern Territory Dept Mines and Energy, Darwin. 158 Specht, R.L. (1988). Vegetation, nutrition and climate. (1). Natural vegetation - ecomorphological characters. Pp. 13-61. In: Mediterranean-type Ecosystems. A Data Source Book. (Ed. R.L. Specht). Kluwer Academic Publ., Dordrecht. 159 Specht, R.L. (1988). Vegetation, nutrition and climate. (4). Climate. Pp. 93-136. In: Mediterranean-type Ecosystems. A Data Source Book. (Ed. R.L. Specht). Kluwer Academic Publ., Dordrecht. 160 Specht, R.L. (1988). Climatic control of ecomorphological characters and species richness in mediterranean eosystems. Pp. 149-155. In: Mediterranean-type Ecosystems. A Data Source Book. (Ed. R.L. Specht). Kluwer Academic Publ., Dordrecht. 161 Stewart, G.R., Hegarty, Elwyn E. and Specht, R.L. (1988). Inorganic nitrogen assimilation in plants of Australian rainforest communities. Physiologia Plantarum 74: 26-33. 162 Specht, R.L. (1988). Geosphere-biosphere interaction in terrestrial ecosystems. Pp. 169-176. In:: Global Change. (Ed. K.D. Cole). Australian Academy of Science, Canberra. 163 Specht, R.L. and Specht, Alison (1989). Species richness of overstorey strata in Australian plant communities. The influence of overstorey growth rates. Australian Journal of Botany 37: 321-336.

59 164 Specht, R.L. and Specht, Alison (1989). Species richness of sclerophyll (heathy) plant communities in Australia. The influence of overstorey cover. Australian Journal of Botany 37: 337-350. 165 Specht, R.L. and Specht, Alison (1989). Canopy structure in Eucalyptus -dominated communities in Australia along climatic gradients. Acta Oecologia, Oecologia Plantarum 10: 191-213. 166 Specht, R.L. and Rundel, P.W. (1990). Sclerophylly and foliar nutrient status of mediterranean-climate plant communities in southern Australia. Australian Journal of Botany 38: 459-474. 167 Specht, R.L. (1990). Changes in the eucalypt forests of Australia as a result of human disturbance. Pp. 177- 197. In: The Earth in Transition. Patterns and Processes of Biotic Impoverishment. (Ed. G. M. Woodwell). Cambridge University Press, New York. 168 Specht, R.L. and Yates, D.J. (1990). Climatic control of structure and phenology of foliage shoots in dicotyledonous overstorey and understorey strata of subtropical plant communities in eastern Australia. Acta Oecologia, Oecologia Plantarum 11: 215-233. 169 Stewart, G. R., Gracia, C.A., Hegarty, Elwyn.E. and Specht, R.L. (1990). Nitrate reductase activity and chlorophyll content in sun leaves of subtropical Australian closed-forest (rainforest) and open-forest communities. Oecologia (Berlin) 82 : 544-551. 170 Specht, R. L., Grundy, Rhonda.I. and Specht, Alison (+ Ruhama Berliner) (1990). Species richness of plant communities — relationship with community growth and structure. Israel Journal of Botany 39 : 465- 480. 171 Batianoff, G.N., Reeves, R.D. and Specht, R.L. (1990). Stackhousia tryonii Bailey: a nickel-accumulating serpentinite-endemic species of central Queensland. Australian Journal of Botany 38 : 121-130. 172 Specht, R.L., Yates, D.J., Sommerville, Jacqui E.M. and Moll, E.J. (1991). Foliage structure and shoot growth in heathlands in the mediterranean-type climate of southern Australia and South Africa. Ecologia Mediterranea 16: 195-207. 173 Specht, R.L. and Clifford, H.T. (1991). Plant invasion and soil seed banks: control by water and nutrients. Pp. 191-204 . In: Biogeography of Mediterranean Invasions. (Eds R.H. Groves and F. di Castri). Cambridge University Press, Cambridge. 174 Specht, R. L., Dettmann, Mary E. and Jarzen, D. M. (1992). Community associations and structure in the Late Cretaceous vegetation of southeast Australasia and Antarctica. Palaeogeography, Palaeoclimatology, Palaeoecology 94 : 283-309. 175 Batianoff, G. N. and Specht, R. L. (1992). Queensland (Australia) serpentine vegetation. Pp. 109 - 128 In: The Vegetation of Ultramafic (Serpentine) Soils. (Eds A. J. M. Baker, J. Proctor and R. D. Reeves) Intercept Publications, Andover, England. 176 Specht, Marion M. and Specht, R. L. (1992). Herbivory of leaves of Banksia oblongifolia, with or without overstorey cover. Pp. 22-27. In: Proceedings of the Sixth International Conference on Mediterranean Ecosystems. Plant-Animal Interactions in Mediterranean-type Ecosystems. (Ed. C. A. Thanos). University of Athens, Greece. 177 Specht, Alison and Specht, R. L. (1993). Species richness and canopy productivity of Australian plant communities. Biodiversity and Conservation 2 : 152 - 167. 178 Specht, Alison and Specht, R. L. (1994). Biodiversity of overstorey trees in relation to canopy productivity and stand density in the climatic gradient from warm temperate to tropical Australia. Biodiversity Letters 2: 39-45. 179 Specht, R. L. (1994). Biodiversity and conservation. Pp. 525-555. In: Australian Vegetation. Second Edition. (Ed. R. H. Groves). Cambridge University Press, Cambridge. 180 Specht, R. L. (1994). Species richness of vascular plants and vertebrates in relation to canopy productivity. Pp. 15-24. In: Plant-Animal Interactions in Mediterranean-Type Ecosystems. (Eds Margarita Arianoutsou and R. H. Groves). Kluwer Scientific Publications, Dordrecht. 181 Specht, Alison and Specht, R. L. (October 1994). The effect of global change on biodiversity (number of species per hectare) in the Mediterranean-type climate of southern Australia. MEDECOS Resúmenes. Noticiero de Biologia, Chile 2(3): 52. 182 Specht, R.L. and Dettmann, Mary E. (1995). Palaeo-ecology of Australia and current physiological functioning of plant communities. Pp. 201-214. In: Time Scales of Biological Responses to Water Constraints: The Case of Mediterranean Biota. (Eds J. Roy, J. Aronson and F. di Castri) SPB Academic Publishing, Amsterdam. 183 Specht, R.L. and Specht, Alison (1995). Global warming: Predicted effects on structure and species richness of Mediterranean ecosystems in southern Australia. Pp. 215-237. In: Time Scales of Biological Responses to Water Constraints: The Case of Mediterranean Biota. (Eds J. Roy, J. Aronson and F. di Castri) SPB Academic Publishing, Amsterdam. 184 Specht, R. L. (1995). El funcionament ecològic de la sabana. Pp. 93-101. In: Biosfera. Vol. 3. Sabanes. (Eds R. Folch et al.) Enciclopèdia Catalana, Barcelona. 185 Specht, R. L. (1996). The influence of soils on the evolution of the eucalypts. Pp. 31-60. In: Nutrition of the Eucalypts. (Eds P. M. Attiwill and M. A. Adams) C.S.I.R.O. Publications, Melbourne. 186 Specht, R.L. (1997). Ecosystem dynamics in coastal dunes of eastern Australia. Pp. 483-495. In: Ecosystems of the World. Vol. 2 C. Dry Coastal Ecosystems. (Ed. E. van der Maarel). Elsevier, Amsterdam. 187 Batianoff, G. N., Reeves, R. D. and Specht, R. L. (1997). The effect of serpentine on vegetation structure, species diversity and endemism in Central Queensland. Pp. 44-50. In:: The Ecology of Ultramafic and Metalliferous Areas. (Eds T. Jaffré, R. D. Reeves and T. Becquer) Centre ORSTOM de Nouméa, New Caledonia. 188 Specht, R. L. and Specht, Alison (1999 Hardback; 2002 Paperback). Australian Plant Communities. Dynamics of Structure, Growth and Biodiversity. Oxford University Press, Melbourne, 492 pp.

60 189 Specht, Alison and Specht, R. L. (2000). Australia: Biodiversity of ecosystems. pp. 307-324. In: Encyclopedia of Biodiversity. (Ed. S. Levin) Academic Press, San Diego, California. 190 Specht, R. L. (2000). Savanna woodland vegetation in the South East District of South Australia: The influence of evaporative aerodynamics on the foliage structure of the understorey invaded by introduced annuals. Austral Ecology 25 588-599. 191 Specht, R. L. (2001). Phosphorus toxicity and pollution: A threat to our Gondwanan heritage. Ecological Management and Restoration 2: 228-230. 192 Specht, R. L. (2002). Phosphate pollution and soil nitrate: Threats to biodiversity in Australia. Pp. 53-70, In: Landscape Health of Queensland. (Eds A. J. Franks, J. Playford and A. Shapcott) Royal Society of Queensland, Brisbane. 193 Martin H. A. & Specht R. L. (2005). Sclerophyll (heathy) understoreys in the Mount Lofty Ranges, South Australia. Transactions of the Royal Society of South Australia 129, 14-24. 194 Specht, R. L., Batianoff, G. N. and Reeves, R. D. (2005). The vegetation continuum on the serpentinite soil catena. pp. 23-40. In: The Rockhampton-Marlborough Area in Marlborough Serpentine, Landscape, Management and Conservation. (Eds A. Exelby, K. Wormington & A. Melzer) Central Queensland University, Rockhampton. 195 Specht, R. L. (Sept. 2005). The International Biological Programme in Australia: Production Processes in Terrestrial Ecosystems. A personal perspective. Bulletin of the. Ecological Society of Australia 35, 2: 13- 14. 196 Specht, R. L., Batianoff, G. N. and Reeves, R. D. (2006). Vegetation structure and biodiversity along the eucalypt forest to rainforest continuum on the serpentinite soil catena in a subhumid area of Central Queensland, Australia. Austral Ecology 31: 394-407. 197 Specht, R. L. and Specht, Alison (2007). Pre-settlement tree density in the eucalypt open-forest on the Brisbane Tuff. Proceedings of the Royal Society of Queensland 113: 9-16. 198 Specht, R. L. (2007). Species richness of rainforest stands on non-serpentinite and serpentinite substrates in the Rockhampton – Marlborough area of Central Queensland. Proceedings of the Royal Society of Queensland 113: 17-35. 199 Specht, R. L. (2009). Structure and species richness in wetland continua on sandy soils in subtropical and tropical Australia. Austral Ecology 34: 761-772. 200 Specht, R. L. (Nov. 2009). Appraising the legacy of the Arnhem Land Expedition: An insider’s perspective. Page 97 In: ‘Barks, Birds and Billabongs’. ‘Exploring the Legacy of the 1948 American-Australian Scientific Expedition to Arnhem Land.’ (Ed. Margo Neale) National Museum of Australia, Canberra. 201 Specht, R. L. and Tyler, M. J. (2010) The species richness of vascular plants and Amphibia in major plant communities in temperate and tropical Australia: relationship with annual biomass production. International Journal of Ecology 2010, Article ID 635852, 17 pages (doi: 10.1155/2010/635852) 202 Specht, R. L. and Specht, Alison (2010). The ratio of foliar nitrogen to foliar phosphorus — A determinant of leaf attributes and height in life-forms of subtropical and tropical plant communities. Australian Journal of Botany 58: 527-538. 203 Specht, R. L. (2011). Development of ecosystem research. International Scholarly Research Network ISRN Ecology, Volume 2011, Article ID 897578, 20 pages. (doi: 10.5402/2011/897578) 204 Specht, R. L. (2012). Plant communities of North Stradbroke Island: development of structure and species richness. Proceedings of the Royal Society of Queensland 117: 181-191. 205 Specht, R. L. (2012). Biodiversity of terrestrial ecosystems in tropical to temperate Australia. International Journal of Ecology 2012, Article ID 35982, 15 pages. (doi:10.1155/2012/359892) 206 Specht, R. L. (2012). American-Australian Scientific Expedition to Arnhem Land (1948) — Its Long-Range Impact. The Open Ecology Journal 2012, Vol. 5: 30 pages. 207 Specht, R. L. (2014). Management of biodiversity in Australia. Macrothink Institute Journal of Environment & Ecology 5: 186-205. 208 Specht, R. L. and Moll, E. J. (2014). Structure and alpha biodiversity of major plant communities in South Africa, a close biogeographical relation with Australia. Macrothink Institute Journal of Environment & Ecology 5: 48-88. 209 Specht, R. L., Montenegro, Gloria and Dettmann, Mary E. (2015). Structure and alpha biodiversity of major plant communities in Chile, a distant biogeographical relation with Australia. Macrothink Institute Journal of Environment & Ecology 6: 21-47. 210 Alison Specht, Matthew P. Bolton, Bryn Kingsford, Raymond L. Specht and Lee Belbin (2018). A story of data won, data lost and data re-found: the realities of ecological data preservation. Biodiversity Data Journal 6: e28073 doi:10.3897/BDJ.6.e28073

(6) Australian plant communities — Historical studies

211 Specht, R. L. (1958). History of botanical exploration in the Arnhem Land Aboriginal Reserve. Pp. 1-8. In: Records of the American-Australian Expedition to Arnhem Land. Vol. 3. Botany and Plant Ecology. (Eds R. L. Specht and C. P. Mountford). Melbourne University Press, Melbourne. 212 Specht, Marion M. and Specht, R. L. (1962). Bibliographia phytosociologica: Australia. Excerpta Botanica, Sectio B 4: 1-58. 213 Specht, Marion M. (1964). The history of zoological collections made in and on the shores of the Arnhem Land Aboriginal Reserve. Pp. 1-7. In: Records of the American-Australian Expedition to Arnhem Land. Vol. 4. Zoology. (Ed R. L. Specht). Melbourne University Press, Melbourne.

61 214 Specht, R. L. (1976). History and contemporary state of terrestrial ecology in Australian universities. The Australian University 14(2): 99-113. 215 Specht, R. L. (1978). Obituary - Desmond Andrew Herbert, C.M.G., D.Sc. Proceedings of the Royal Society of Queensland 89: 134-135. 216 Specht, R. L. (1981). Developments in terrestrial ecology in Australia. Pp. 387-415. In: Handbook of Contemporary Developments in World Ecology. (Eds E. J. Kormondy and J. F. McCormick). Greenwood Press, Westport, Conn. 217 Specht, R. L. and Specht, Marion M. (1995). The 'Web of Life' in Queensland. (Australian Academy of Science) Proceedings of the Royal Society of Queensland 105: 55-60. 218 Specht, R. L. and Specht, Marion M. (1996). History of identification of Queensland plant communities — from Leichhardt (1844-45) to 1995. Proceedings of the Royal Society of Queensland 106: 17-26. 219 Specht, R. L. and Specht, Alison (Eds) (1998). Personal Experiences on the 'Coast of Adventure'. American- Australian Scientific Expedition to Arnhem Land, March-November 1948. Arnhem Land Reunion, Sydney, 27-28th June 1998. Southern Cross University, Lismore, N.S.W. 220 Specht, R. L. (2002). Wood, Joseph Garnet (1900 – 1959). P. In: Australian Dictionary of Biography (Ed. J. Ritchie) Canberra. 221 Specht, R. L. (2002). Obituary: Margaret McArthur Oliver 1919 – 2002. Australian Aboriginal Studies 2002/2: 122-124. Asia Pacific Journal of Clinical Nutrition 12: 127-128. 222 Specht, R. L. (2004). Obituary. Brian Price Billington 24 February 1923 – 6 August 2004. Australian Aboriginal Studies 2004/2: 133. 223 Specht, R. L. and McCarthy, F. D. (2005). The snake woman story’ of Hemple Bay, Groote Eylandt: An Enindilyakwa legend of a tsunami in the Gulf of Carpentaria. Proceedings of the Royal Society of Queensland 112: 77-82. 224 Robertson, D. J., Specht, R. L. and Nurcombe, B. (2014). Assessment of mental health in Indigenous Australians: the role of John Ewart Cawte, a pioneer in transcultural research. Health & History 16: 107- 114.

(7) Australian plant communities — Ethnobotany

225 Specht, R. L. (1958). An introduction to the ethno-botany of Arnhem Land. Pp. 479-504. In: Records of the American-Australian Expedition to Arnhem Land. Vol. 3. Botany and Plant Ecology. (Eds R. L. Specht and C. P. Mountford). Melbourne University Press, Melbourne. 226 Wahlqvist, M. L. and Specht, R. L. (1998). Food variety and biodiversity: Econutrition. Asia Pacific Journal of Clinical Nuttrition 7: 314-319. 227 McArthur, M.argaret, McCarthy, F. D. and Specht, R. L. (2000). Nutrition studies (1948) of nomadic Aborigines in Arnhem Land, northern Australia. Asia Pacific Journal of Clinical Nuttrition 9: 215-223. 228 McCarthy, F. D. and McArthur, M. (1960). The food quest and the time factor in aboriginal economic life. pp. 145-194. In: Records of the American-Australian Expedition to Arnhem Land. Vol. 2. Anthropology and Nutrition. (Ed. C. P. Mountford). Melbourne University Press, Melbourne. 229 Specht, R. L. and Specht, Alison (1999). Aboriginal impact. Pp. 132-156. In: Australian Plant Communities. Dynamics of Structure, Growth and Biodiversity. Oxford University Press, Melbourne. (R. L. Specht, as botanist on the 1948 Arnhem Land Expedition, was closely involved with all aboriginal nutrition studies).

(8) Reviews by R. L. Specht

Aston, Helen I. (1973). Aquatic Plants of Australia. Melbourne University Press. — Search 5: 401 (1974). Beadle, N. C. W. (1981). The Vegetation of Australia. Cambridge University Press, Cambridge. — Journal of Ecology 70: 905-906 (1982). Carr, D. J. and Carr, S. G. Maisie (Eds) (1981). People and Plants in Australia. Academic Press, Sydney. — Search 13: 258 (1982). Carr, D. J. and Carr, S. G. Maisie (Eds) (1981). Plants and Man in Australia. Academic Press, Sydney. — Search 13: 258 (1982). Sen, D. N. and Rajpurohit, K. S. (Eds) (1982). Tasks of Vegetation Science. Vol. 2. Contributions to the Ecology of Halophytes. Junk, The Hague. — J. Aust. Inst. Agric. Sci. 48: 232-233 (1982).

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