OFR 151.pdf - CRC LEME
OFR 151.pdf - CRC LEME OFR 151.pdf - CRC LEME
SECTION 2 (THE NATURE OF FOSSIL EVIDENCE) Fossils are the traces of once-living organisms that have been buried by natural processes and subsequently preserved in whole or in part. This definition covers skeletal and cell wall material of any size, whether chemically unaltered, reduced to mineral carbon, replaced by other minerals such as silica, calcite, limonite and pyrite or reduced to impressions (casts and moulds), as well as excreted material, and tracks, trails and borings. Microanalytical and geochemical techniques allow some fossil organisms to be identified from organic residues (organic trace fossils). Examples include marine algae such as acritarchs and dinoflagellates, and C4 grasses, which possess distinctive biochemical ‘signatures’. As an adjective, ‘fossil’ is used more widely, to cover any entity of perceived geological antiquity whether or not the object is extant. Examples include fossil ice wedges (sedimentary casts of ice wedges) and ‘living fossils’ such as Wollemi Pine (Macphail et al. 1995). 2.1 Taphonomy Taphonomy encompasses the post-mortem history of the organisms. Taphonomic processes include the decomposition phase after death (necrophysis), the sedimentological history of fossil remains (biostratinomy), and chemical and physical changes in the fossil between burial and collection (diagenesis). 2.1.1 Fossil assemblages With few exceptions, the individual accumulations of fossils (assemblages) represent geologically instantaneous records of past life. Stratified sequences, which cover much longer periods of geological time, may provide a time series record of past life. The value of a fossil assemblage as proxy-climatic evidence depends on the degree to which the individual fossils can be associated directly to past environments or indirectly via living relatives whose ecological relationships are known. Preservation and geological age are important constraints. In most instances, the palaeoclimatic inferences are qualitative, e.g. warmer and wetter or, if quantitative, are expressed as a range (climatic envelope). Geologic evidence such as glendonites and the ratios of naturally occurring isotopes can provide more or less precise quantitative evidence of past climates. 2.1.2 Palaeobotany Palaeobotany is the study of plant remains. The discipline may or may not be distinguished from palaeontology, which can encompass animal remains only (colloquial usage) or all fossils (dictionary definition/industry usage) according to context. Palaeobotanical evidence is subdivided into two main classes according to the size of the remains: macrofossils (chiefly stems, foliage, flowers and fruits) and microfossils (chiefly algal cysts, spores and pollen grains). The two forms of evidence are complementary in that they usually reflect different elements in the palaeovegetation. It is important to note that good preservation of plant macrofossils is no guarantee that microfossils of the same plants will have been preserved or can be identified to the same taxonomic level (and vice versa). 43
2.1.3 Palaeoecology Palaeoecology is the study of the interactions of organisms with one another and with their environment in the past. It differs from the ecology of living organisms in that incomplete preservation usually prevents direct observation of many aspects of the biota. 2.2 Plant macrofossils Plant macrofossil assemblages (macrofloras) constitute a highly detailed record of past vegetation that is strongly biased towards plants growing close to water, e.g. on the banks of sluggish rivers or around lakes (Burnham 1989, Christophel and Greenwood 1987a, Greenwood 1992, 1994, Alexander et al. 1999). 2.2.1 Taphonomic constraints Plant remains are subjected to a number of processes between the time of abscission and their burial in sediments where conditions may or may not lead to long-term preservation. For example, cellulose and lignin, which are the major compounds making up cell walls, are the food source for many fungi and soil invertebrates. Similarly, the size of many plant remains makes them susceptible to physical attrition during transport and deposition, especially by water and wind. Accordingly, except under anoxic conditions, only the more robust plant parts such as roots, stems, twigs and leaves are preserved in an unaltered state. The most commonly found remains are leaves, particularly those of species with naturally dehiscent foliage such as deciduous trees. 2.2.2 Taxonomic constraints The taxonomic level to which plant macrofossils can be identified using foliage or wood is high, often to genus or species level, since many of the taxonomic characters are the same as the characters used to identify modern plant species or genera. For example, leaf cuticle is chemically stable, and microscopic features such as the distribution of stomata and presence of trichomes (plant hairs) allow small fragments to be compared with those of living species with great accuracy, even when primary taxonomic evidence such as flowers and fruits are absent. Similarly the arrangement of the various types of cells making up wood (xylem) allows some stem remains to be assigned to modern genera. If such characters are not preserved, e.g. in impressions and casts, it can be difficult or impossible to determine phylogenetic affinities accurately (Jordan and Hill 1999). The resistance to decay of wood and leaf cuticle is similar to that of fossil spores and pollen and both are major components of the finely disseminated (acid-resistant) detritus (kerogen) preserved in sedimentary rocks (cf. Rowett 1993a). 2.3 Plant microfossils Fossil pollen and spores (miospores) together with phytoliths and algal cysts are by far the most abundant and widespread of all plant remains. Although dispersed by much the same transport processes, the distinction between spores and pollen is an important one in plant migration, and therefore how the palaeobotanical record mirrors past climates: • Spores are produced by fungi, mosses, liverworts, fern allies and ferns (collectively termed cryptogams) and are functionally equivalent to the seeds of flowering plants in that they germinate to produce the next generation of plants. Dispersal is by wind and/or water, less commonly by foraging insects. 44
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2.1.3 Palaeoecology<br />
Palaeoecology is the study of the interactions of organisms with one another and with their<br />
environment in the past. It differs from the ecology of living organisms in that incomplete<br />
preservation usually prevents direct observation of many aspects of the biota.<br />
2.2 Plant macrofossils<br />
Plant macrofossil assemblages (macrofloras) constitute a highly detailed record of past<br />
vegetation that is strongly biased towards plants growing close to water, e.g. on the banks of<br />
sluggish rivers or around lakes (Burnham 1989, Christophel and Greenwood 1987a,<br />
Greenwood 1992, 1994, Alexander et al. 1999).<br />
2.2.1 Taphonomic constraints<br />
Plant remains are subjected to a number of processes between the time of abscission and their<br />
burial in sediments where conditions may or may not lead to long-term preservation. For<br />
example, cellulose and lignin, which are the major compounds making up cell walls, are the<br />
food source for many fungi and soil invertebrates. Similarly, the size of many plant remains<br />
makes them susceptible to physical attrition during transport and deposition, especially by<br />
water and wind. Accordingly, except under anoxic conditions, only the more robust plant<br />
parts such as roots, stems, twigs and leaves are preserved in an unaltered state. The most<br />
commonly found remains are leaves, particularly those of species with naturally dehiscent<br />
foliage such as deciduous trees.<br />
2.2.2 Taxonomic constraints<br />
The taxonomic level to which plant macrofossils can be identified using foliage or wood is<br />
high, often to genus or species level, since many of the taxonomic characters are the same as<br />
the characters used to identify modern plant species or genera. For example, leaf cuticle is<br />
chemically stable, and microscopic features such as the distribution of stomata and presence<br />
of trichomes (plant hairs) allow small fragments to be compared with those of living species<br />
with great accuracy, even when primary taxonomic evidence such as flowers and fruits are<br />
absent. Similarly the arrangement of the various types of cells making up wood (xylem)<br />
allows some stem remains to be assigned to modern genera. If such characters are not<br />
preserved, e.g. in impressions and casts, it can be difficult or impossible to determine<br />
phylogenetic affinities accurately (Jordan and Hill 1999).<br />
The resistance to decay of wood and leaf cuticle is similar to that of fossil spores and pollen<br />
and both are major components of the finely disseminated (acid-resistant) detritus (kerogen)<br />
preserved in sedimentary rocks (cf. Rowett 1993a).<br />
2.3 Plant microfossils<br />
Fossil pollen and spores (miospores) together with phytoliths and algal cysts are by far the<br />
most abundant and widespread of all plant remains. Although dispersed by much the same<br />
transport processes, the distinction between spores and pollen is an important one in plant<br />
migration, and therefore how the palaeobotanical record mirrors past climates:<br />
• Spores are produced by fungi, mosses, liverworts, fern allies and ferns (collectively<br />
termed cryptogams) and are functionally equivalent to the seeds of flowering plants in<br />
that they germinate to produce the next generation of plants. Dispersal is by wind<br />
and/or water, less commonly by foraging insects.<br />
44