OFR 151.pdf - CRC LEME

OFR 151.pdf - CRC LEME OFR 151.pdf - CRC LEME

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The 10 μm sieved, oxidised extract minimises the time required to determine the age of the sample (using larger palynomorphs) whilst the other extracts provide the quantitative information required to make reliable estimates of the palaeovegetation and climate. 8.2.4 Improved age control Only rarely can fossil assembles in inland Australia be directly dated using geochronometric techniques. At present, macrofloras and some local faunas are directly or indirectly dated via criteria established for basins hundreds to thousands of kilometres away. This review indicates that fossil pollen and spores provide the only cheap, moderately reliable means of dating terrestrial fossil floras and faunas across the range of depositional environments found in Australia. The existing data make it highly likely that local palynostratigraphies can be synthesised from disparate short sequences and spot samples for the various regions in inland southern, central and northern Australia. An alternative is to use systematic changes in regional climate as a basis for developing eco-stratigraphic criteria to date continental fossil assemblages. 8.2.5 Application of Quaternary quantitative (objective) analytical techniques Three of the objective (statistical) techniques used by to capture palaeoclimatic information for the Late Quaternary are likely to be applicable to Cretaceous and Tertiary microfloras. These are Isopollen mapping, Principal components analysis and Biome analysis respectively. 1. Isopollen maps Isopollen mapping was developed in North West Europe and North America as a way of visually tracking the climatically forced expansion of individual species and vegetation types during the Late Pleistocene and Holocene (Birks and Saarnisto 1975, Bernabo and Webb 1977, Birks and Birks 1980). The method is analogous to the preparation of weather maps in that lines linking sites with similar relative pollen values can be used to reconstruct patterns in past vegetation and climate. An example is the 1998 video clip (SiteSeer) showing the postglacial expansion of the major tree taxa across North America (http://www.ngdc.gov/paleo/siteseer.html). Essential requirements are statistically reliable quantitative data based on a minimum of 200- 250 counts, good site coverage across the region of interest; and reliable age control. On present indications, these conditions are best met for the Early Cretaceous where sites preserving diverse microfloras can be found across much of the continent. For example, quantitative analysis of ~1000 microfloras (about 2 years work using processed material housed at the various State geological surveys) would allow isopollen maps to be prepared for all palynostratigraphic subdivisions of Berriasian to Albian time. By comparing isopollen maps of commonly occurring Cretaceous taxa against, for example, the changing position of Cretaceous shorelines, it should be possible to objectively infer palaeoecological relationships. These in turn provide a method for tracking climatic developments during the Early Cretaceous. 119

2. Principal components analysis (PCA) This technique is widely used in ecological and palaeoecological research to identify groups of associated or correlated taxa (Birks and Birks 1980, Birks and Gordon 1985). The appropriate routine is included in statistical software used to generate Late Quaternary pollen histograms, for example Tilia. PCA can be useful in the reconstruction of both Cretaceous and Tertiary palaeoclimates in two ways. Firstly, it can help link taxa whose ecology is uncertain with taxa whose climatic preferences have been established by other means. Secondly, it can help identify areas with similar bioclimates by identifying recurrent groups of taxa. 3. Biome analysis The biomization method was developed to resolve problems arising from direct comparisons of Holocene and Quaternary palaeoecological data from very widely separated sites. Rather than focus on shared taxa (if any), the technique utilises the well-known phenomenon of functional convergence in the ecophysiology of unrelated plants growing in similar physical environments. This convergence may be expressed at the molecular or whole plant level. Examples are the evolution of the C4 photosynthetic pathway in unrelated plant families occupying drier, warmer habitats and the evolution of a drip-tip on the leaves of many unrelated rainforest angiosperms. The method involves: (1) assigning plants to plant functional types (PFTs); (2) defining biomes in terms of the PFTs of the constituent taxa; and (3) using observed bioclimatic relationships to reconstruct patterns of climate at a given time in the past. Mathematically, the method involves calculating the affinity (affinity score) of each fossil flora to each biome defined by PFTs and assigning the flora to the biome (and climate) with which it has the highest affinity (references in Prentice and Webb 1998). The formula used for Holocene microfloras combines presence/absence and relative abundance data weighted by threshold values, which take account of the different representation of the parent plants by pollen or spores. Prentice and Webb (1998) discuss how the technique can be applied to the mid Holocene. Horrell (1991) has used an analogous approach to reconstruct global climatic zones for the Maastrichtian. Holocene floras are a special case in that almost all microfossil taxa can be assigned to plant functional types in the certain knowledge that the fossil plants possessed the same ecophysiological adaptations as living representatives. Only ~200 of the >300 described Tertiary fossil species have presumed NLRs whilst many more undescribed taxa have not been systematically recorded because of small size, simple morphologies and lack of (known) biostratigraphic value. Nevertheless sites that preserve both plant macrofossils and microfossils confirm the general applicability of modern relationships (phylogenetic and ecophysiological) to Tertiary situations (Macphail et al. 1994), even if this may not be the case for the earlier Cretaceous floras. Accordingly, with appropriate resources it would seem possible to develop a mathematical model along the lines of Prentice and Webb's (1998) formula, which could assign Tertiary and possibly Late Cretaceous floras to biomes defined by PFTs. Logically, any formal quantitative analysis would be empirical and begin with either the Pliocene microfloras, because of the significant number of taxa with moderately well known biome relationships, or the Oligo-Miocene macrofloras, because their ecophysiological adaptations to primary forcing factors such as humidity can be directly observed. The method reduces often very large numbers of fossil taxa into more manageable ecophysiological types using the basic physiognomic characteristics of life-form, leaf-form and 120

2. Principal components analysis (PCA)<br />

This technique is widely used in ecological and palaeoecological research to identify groups<br />

of associated or correlated taxa (Birks and Birks 1980, Birks and Gordon 1985). The<br />

appropriate routine is included in statistical software used to generate Late Quaternary pollen<br />

histograms, for example Tilia.<br />

PCA can be useful in the reconstruction of both Cretaceous and Tertiary palaeoclimates in<br />

two ways. Firstly, it can help link taxa whose ecology is uncertain with taxa whose climatic<br />

preferences have been established by other means. Secondly, it can help identify areas with<br />

similar bioclimates by identifying recurrent groups of taxa.<br />

3. Biome analysis<br />

The biomization method was developed to resolve problems arising from direct comparisons<br />

of Holocene and Quaternary palaeoecological data from very widely separated sites. Rather<br />

than focus on shared taxa (if any), the technique utilises the well-known phenomenon of<br />

functional convergence in the ecophysiology of unrelated plants growing in similar physical<br />

environments. This convergence may be expressed at the molecular or whole plant level.<br />

Examples are the evolution of the C4 photosynthetic pathway in unrelated plant families<br />

occupying drier, warmer habitats and the evolution of a drip-tip on the leaves of many<br />

unrelated rainforest angiosperms.<br />

The method involves: (1) assigning plants to plant functional types (PFTs); (2) defining<br />

biomes in terms of the PFTs of the constituent taxa; and (3) using observed bioclimatic<br />

relationships to reconstruct patterns of climate at a given time in the past. Mathematically,<br />

the method involves calculating the affinity (affinity score) of each fossil flora to each biome<br />

defined by PFTs and assigning the flora to the biome (and climate) with which it has the<br />

highest affinity (references in Prentice and Webb 1998). The formula used for Holocene<br />

microfloras combines presence/absence and relative abundance data weighted by threshold<br />

values, which take account of the different representation of the parent plants by pollen or<br />

spores. Prentice and Webb (1998) discuss how the technique can be applied to the mid<br />

Holocene. Horrell (1991) has used an analogous approach to reconstruct global climatic<br />

zones for the Maastrichtian.<br />

Holocene floras are a special case in that almost all microfossil taxa can be assigned to plant<br />

functional types in the certain knowledge that the fossil plants possessed the same<br />

ecophysiological adaptations as living representatives. Only ~200 of the >300 described<br />

Tertiary fossil species have presumed NLRs whilst many more undescribed taxa have not<br />

been systematically recorded because of small size, simple morphologies and lack of (known)<br />

biostratigraphic value. Nevertheless sites that preserve both plant macrofossils and<br />

microfossils confirm the general applicability of modern relationships (phylogenetic and<br />

ecophysiological) to Tertiary situations (Macphail et al. 1994), even if this may not be the<br />

case for the earlier Cretaceous floras. Accordingly, with appropriate resources it would seem<br />

possible to develop a mathematical model along the lines of Prentice and Webb's (1998)<br />

formula, which could assign Tertiary and possibly Late Cretaceous floras to biomes defined<br />

by PFTs. Logically, any formal quantitative analysis would be empirical and begin with<br />

either the Pliocene microfloras, because of the significant number of taxa with moderately<br />

well known biome relationships, or the Oligo-Miocene macrofloras, because their<br />

ecophysiological adaptations to primary forcing factors such as humidity can be directly<br />

observed.<br />

The method reduces often very large numbers of fossil taxa into more manageable ecophysiological<br />

types using the basic physiognomic characteristics of life-form, leaf-form and<br />

120

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