A reconstruction of Archean biological diversity based on molecular fossils from the 2.78 to 2.45 billion-year-old Mount Bruce Supergroup, Hamersley Basin, Western Australia
Introduction
Microfossils, stromatolites and isotopes of sedimentary carbon and sulfur all indicate that microorganisms inhabited Earth during the Archean, the time before 2.5 billion years ago (Ga) (Fig. 1). The fossils and isotopes also provide circumstantial evidence for the early evolution of some physiological attributes and metabolic pathways. Sedimentary organic matter strongly depleted in the carbon isotope 13C implies the presence of archaeal methanogens and bacterial methanotrophs (Hayes, 1983), fractionated sulfide isotopes in sulfate crystals suggest the existence of mesophilic bacterial sulfate-reducers (Shen et al., 2001), and palimpsest filament tufts in lacustrine stromatolites are consistent with phototropic cyanobacteria (Buick, 1992). However, these lines of evidence depend on chains of inference with concomitant compounding uncertainties. The oldest fossils well enough preserved to be recognized as a certain member of an extant clade only appear in the mid-Paleoproterozoic, providing diagnostic and unequivocal evidence for organisms of the phylum Cyanobacteria at 2.15 Ga (Hofmann, 1976). The oldest remains of possible eukaryotes were discovered in iron deposits 1.87 Ga old Han and Runnegar 1992, Schneider et al 2002, while morphologic evidence for the domain Archaea is not known from any part of the fossil record. Therefore, the phylogenetic position of life in the Archean remains largely obscure. Potentially, a better insight into early Precambrian microbial diversity can be obtained from molecular fossils (biomarkers) preserved in sedimentary rocks.
This paper discusses the paleobiological and paleoenvironmental significance of biomarkers detected in late Archean sedimentary rocks from the Pilbara region of Western Australia Brocks et al 1999, Brocks et al 2003. Biomarkers, their distribution in the Archean samples, their occurrence in other Precambrian strata and their potential biologic source will be discussed according to hydrocarbon class in the following order: normal and branched alkanes, cyclohexylalkanes, adamantanes, tricyclic terpanes, hopanes and the diverse range of steranes and steroids.
A companion paper in this issue (Brocks et al., 2003) gives a detailed assessment of syngeneity of the biomarkers and addressing the possibility that the Archean host rocks were contaminated during drilling and storage, or were adulterated by migrating fluids during post-Archean history. It is concluded that adamantanes and polyaromatic hydrocarbons of the Hamersley Group are ‘certainly syngenetic’, and aliphatic hydrocarbons and polycyclic biomarkers of the Hamersley and Fortescue Group ‘probably syngenetic’. Therefore, the age of the molecules that contain the biologic information is not fully resolved. Hence, paleobiological interpretation presented in this paper should be cited cautiously and with reference to the remaining uncertainty of syngeneity (Brocks et al., 2003).
The interpretation of Archean and Proterozoic biomarkers is complicated by two factors. One is the fragmentary knowledge of biomarker distributions across the great diversity of extant organisms and the second is the problem of how reasonable it is to extrapolate modern biomarker relationships back in time over several billion years. The biological interpretation of geolipids is almost exclusively based on the distribution of biolipids in extant organisms. However, the full biolipid repertoire is only known for a small percentage of microorganisms that have been cultured (Volkman et al., 1993). It is possible, therefore, that many lipids might have a broader biological distribution and more or less taxonomic value than currently known. More problematic, however, are the uncertainties associated with attributing biological sources to biomarkers from rocks several hundred million to billions of years old. Particular pathways of lipid biosynthesis, such as the modification of sterol side-chains, might have evolved independently in different lineages or could have been passed by lateral gene transfer between lineages. So lipids believed to be diagnostic of extant taxonomic groups could also have been prevalent in extinct clades.
Section snippets
N-alkanes
By far the most abundant hydrocarbons in all late Archean bitumens are n-alkanes (Brocks et al., 2003). Homologues with chain lengths from 9 to 23 carbon atoms and in one case 28 carbon atoms were detected under GC-FID (gas chromatography–flame ionization detection) conditions. The n-alkane distribution is commonly unimodal with a strong predominance of low-molecular weight homologues, a composition generally controlled by the high thermal maturity of the samples. Odd or even predominance of
Reconstruction of late archean biodiversity
To reconstruct the late Archean biosphere, biomarker data should be viewed in the context of current paleontological knowledge. The following section reviews fossil and isotope evidence for the antiquity of the three domains of life—Bacteria, Archaea and Eucarya—and discusses new knowledge provided by Archean biomarkers (Table 1).
Acknowledgements
Supported by the American Chemical Society Petroleum Research Fund (R.B.), Geoscience Australia (J.J.B.) and a School of Geosciences, University of Sydney special stipend (J.J.B.). The authors thank Janet Hope, Carolyn Sandison, Christian Thun, Natalie Johns, Dan Gardner and Ian Atkinson for expert technical assistance, Julie Kamprad and Algis Juodvalkis for XRD measurements, and Emmanuelle J. Javaux and Andrew H. Knoll for insightful discussions on early eukaryote evolution. We thank Rio Tinto
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