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View Open Access the Structure of Biodiversity – Insights from Molecular Phylogeography Godfrey M Hewitt* Frontiers in Zoology BioMed Central Review Open Access The structure of biodiversity – insights from molecular phylogeography Godfrey M Hewitt* Address: Biological Sciences, UEA, Norwich NR4 7TJ, UK Email: Godfrey M Hewitt* - [email protected] * Corresponding author Published: 26 October 2004 Received: 18 October 2004 Accepted: 26 October 2004 Frontiers in Zoology 2004, 1:4 doi:10.1186/1742-9994-1-4 This article is available from: http://www.frontiersinzoology.com/content/1/1/4 © 2004 Hewitt; licensee BioMed Central Ltd. This is an open-access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/2.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. Abstract DNA techniques, analytical methods and palaeoclimatic studies are greatly advancing our knowledge of the global distribution of genetic diversity, and how it evolved. Such phylogeographic studies are reviewed from Arctic, Temperate and Tropical regions, seeking commonalities of cause in the resulting genetic patterns. The genetic diversity is differently patterned within and among regions and biomes, and is related to their histories of climatic changes. This has major implications for conservation science. Introduction chloroplast (cp) in plants and non-coding nuclear (nc) Phylogeography, named by Avise et al in 1987 [1], is a regions in both animals and plants. MtDNA has a rela- recent and rapidly developing field that concerns the geo- tively fast rate of nucleotide divergence, well suited to graphical distribution of genealogical lineages. It grew examining events over the last few million years, but those from the newly acquired technical ability to obtain DNA of cpDNA and ncDNA are an order of magnitude lower sequence variation from individuals across a species and consequently less useful for such young divergences. range, and from this to reconstruct phylogenies. These are More slowly evolving sequences are required for deeper then plotted geographically to display their spatial rela- phylogenetic history. For more recent events, like the last tionships and deduce the evolutionary origins and history 10 thousand years, highly variable markers are needed, of populations, subspecies and species [2,3]. Genetic rela- such as microsatellites and AFLPs, but whilst useful for tionships between species based on polytene chromo- population studies they suffer from homoplasy and pro- some banding patterns had been used earlier to deduce duce equivocal genealogies [3]. the geographic history of colonization and speciation by Drosophila of the Hawaiian Islands [4], and allozyme var- Techniques for obtaining DNA sequence information are iation may still complement DNA information, but the still advancing rapidly, with whole genome sequences ready access to mitochondrial (mt) DNA sequences being produced in a growing number of organisms. This opened the door to most animal species and generated allows sequences and markers to be identified and devel- this new field [5]. oped for many types of investigation, and some will be useful for phylogeographic studies. In particular, it is clear DNA methods that genealogical data is required from several independ- Whilst mtDNA has lead the way in animal phylogeogra- ent nuclear loci to provide a fuller and more reliable his- phy, other DNA sequences are used, most commonly tory of the species [6]. Single nucleotide polymorphisms Page 1 of 16 (page number not for citation purposes) Frontiers in Zoology 2004, 1:4 http://www.frontiersinzoology.com/content/1/1/4 (SNPs) are also becoming available across the genome, Paleoclimate and Paleobiology which will produce comprehensive measures of genetic The very different field of paleoclimatology is also experi- diversity and allow the construction of better population encing great advances. The results of these are most perti- histories [7]. nent to phylogeographic explanations, since they reveal the past environmental conditions and changes that have Analytical approaches molded the evolutionary processes producing the present The advance in DNA technology is producing a wealth of genetic structure. They provide a framework in which the data for individuals, populations and species, and there phylogeny may be reconstructed. Past conditions can be are concomitant developments in analytical methods to deduced from carbon and oxygen isotopes, radiolarian divine demographic history and evolutionary relation- skeletons, pollen grains and other residues from the sea ships, and to test their significance. This progress in anal- bed, lake bottoms and ice sheets [16]. Novel information ysis is facilitated by access to increasingly powerful sources like insect exoskeletons, coral terraces and stalag- desktop computers on which the increasingly sophisti- mites are adding to this [17-20]. Such records show that cated software can be used. Haplotype sequences of a par- earth's climate has been cooling for some 60 my with peri- ticular DNA region can be ordered into a genealogical tree odic (21, 41 and 100 kyr) global oscillations producing or network, and hence produce their phylogeny. When increasingly severe ice ages through the Quaternary (2.4 combined with their population frequency and geo- my). These involved greatly enlarged ice sheets and sur- graphic distribution, this provides a strong basis for infer- rounding permafrost, and the lower temperatures and ences on the evolutionary history of the populations and reduced water availability caused great changes in the dis- species. The usual phylogeographic approach is to build a tribution of species as demonstrated by the fossil record phylogeny from haplotype sequences using distance, par- [16,21]. Nested within the major 100 kyr cycles are mil- simony and maximum likelihood methods and then rep- lennial scale oscillations, which can occur rapidly and are resent the lineages geographically. There are several often severe [22,23]. Changes of 7–15°C may occur in approaches that are regularly used, such as DNA Distance decades and persist for centuries, as happened most Phylogeography, Nested Clade Analysis, Haplotype Net- recently in the Younger Dryas (11 kya), where fossils works, Sequence Mismatch Distribution and Genetic and record shifts in the distributions of species. Demographic Simulation, e.g. PAUP and GeoDis [8-10]. This last approach uses computers to explore broadly how These major changes in distributions of species occurred DNA markers evolve in specified molecular, spatial and latitudinally as the ice sheets advanced and retreated, alti- demographic conditions over history, and is being used tudinally in major mountain regions, and also longitudi- increasingly [11]. Recent developments seek to use the nally where new dispersal routes became available, as for genetic data to estimate the demographic history of a pop- example the Bering land bridge produced by the lowered ulation, the dates of historical bottlenecks or expansions, sea level. The demographic fluctuations and adaptive the size of ancestral populations, the location of refugial challenges produced by such range changes would have areas, the dates of divergence, the extent of migration and had both stochastic and selective effects on the genetic gene flow, the extent of fragmentation, and the sequence variation and architecture, and the consequences of these of such events to produce the present geographic distribu- can be studied by genetic and phylogeographic tion of genotypes, e.g. [12-15]. We can expect further approaches. Thus the once distinct fields of paleobiology developments to provide even more discriminating and phylogeography are now being combined, and have analyses. much to tell us about how present biodiversity was structured. Each DNA sequence has its own genealogy and they may evolve at different rates. Furthermore, the various meth- Fossil and Genetic Signals of Range Changes ods of analysis probe different aspects of the molecular With some effort it is now possible to obtain DNA data and spatial history. Consequently, to reconstruct a species from specimens across the present range of a species, but phylogeographic history one would ideally like to use a fossil data is often more limited or absent. The most use- range of sequences (including nuclear, cytoplasmic, sex- ful fossil data are for Europe and North America, which linked, autosomal, conserved, neutral, high and low have extensive networks of pollen cores; a few span 3 ice mutation rate) and apply a suite of pertinent analyses. ages (400 kyr), several reach the last interglacial (125 kyr), This is not easy and often not possible with resources and a larger number cover back to the last glacial maxi- available. However, technological advances for molecules mum (LGM 23-18 kyr) [21]. There are also some helpful and computers have been explosive in the past decade, detailed series of beetle exoskeletons [24]; animal bones making much more detailed analysis possible today than and plant macrofossils tend to be localized and discontin- only a few years ago, and this looks set to continue. uous, but are nonetheless useful markers of time and place. Reconstructions of paleovegetation have been Page 2 of 16 (page number not for citation purposes) Frontiers in Zoology 2004, 1:4 http://www.frontiersinzoology.com/content/1/1/4 made, e.g. [25], which are quite detailed from the LGM to lations and processes involved in colonization [30]; this the present, and when
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