Theodosius Dobzhansky once remarked that nothing in biology makes sense other than in the light of evolution, thereby emphasising the central role of evolutionary studies in providing the theoretical context for all of biology. It is perhaps surprising then that evolutionary biology has played such a small role to date in Antarctic science. This is particularly so when it is recognised that the polar regions provide us with an unrivalled laboratory within which to undertake evolutionary studies. The Antarctic exhibits one of the classic examples of a resistance adaptation (antifreeze peptides and glycopeptides, first described from Antarctic fish), and provides textbook examples of adaptive radiations (for example amphipod crustaceans and notothenioid fish). The land is still largely in the grip of major glaciation, and the once rich terrestrial floras and faunas of Cenozoic Gondwana are now highly depauperate and confined to relatively small patches of habitat, often extremely isolated from other such patches. Unlike the Arctic, where organisms are returning to newly deglaciated land from refugia on the continental landmasses to the south, recolonization of Antarctica has had to take place by the dispersal of propagules over vast distances. Antarctica thus offers an insight into the evolutionary responses of terrestrial floras and faunas to extreme climatic change unrivalled in the world. The sea forms a strong contrast to the land in that here the impact of climate appears to have been less severe, at least in as much as few elements of the fauna show convincing signs of having been completely eradicated.

Published and unpublished data are used to investigate possible mechanisms of species diversification in two key groups of Antarctic organisms: the Notothenioidei and the Euphausiidae. Species distributions are mapped onto molecular phylogenies, and this evidence is interpreted in light of the various ecological and historical factors which characterize the Southern Ocean. The joint effect of diverse agents (vicariance, “jump” dispersal) appears to be determinant in several cases for species divergence. A review of results from population genetics studies, together with new molecular evidence, confirm the importance of physical barriers (oceanographic fronts) in reducing migration, thereby promoting speciation, at least in notothenioids.

A short summary of some the most important hypotheses on the evolution of Southern Ocean peracarid crustaceans and some of the potential reasons for the high biodiversity of this taxon is presented. Besides the knowledge of horizontal and vertical distribution patterns of Southern Ocean peracarids, the importance of the evolution of the notothenioid fishes on the evolution of the Peracarida is discussed. Key questions for a better understanding of evolutionary biology of Southern Ocean Peracarida are highlighted.

Antarctica is a continental island and the waters of its shelf and upper slope are an insular evolutionary site. The shelf waters resemble a closed basin in the Southern Ocean, separated from other continents by distance, current patterns and subzero temperatures. The benthic fish fauna of the shelf and upper slope of the Antarctic Region includes 213 species with higher taxonomic diversity confined to 18 families. Ninety-six notothenioids, 67 liparids and 23 zoarcids comprise 45%, 32% and 11% of the fauna, a combined total of 88%. In high latitude (71–78°S) shelf areas notothenioids dominate abundance and biomass at levels of 90–95%. Notothenioids are also morphologically and ecologically diverse. Although they lack a swim bladder, the hallmark of the notothenioid radiation has been repeated diversification into water column habitats. There are pelagic, semipelagic, cryopelagic and epibenthic species. Notothenioids exhibit the disproportionate speciosity and high endemism characteristic of fish species flock. Antifreeze glycopeptides originating from a transformed trypsinogen gene are a key innovation. It is not known when the modern Antarctic shelf fauna assumed its current taxonomic composition. A late Eocene fossil fauna was taxonomically diverse and cosmopolitan. There was a subsequent faunal replacement with little carryover of families into the modern fauna. Basal notothenioid clades probably diverged in Gondwanan shelf locations during the early Tertiary. Dates inferred from molecular sequences suggest that phyletically derived Antarctic clades arose 15–5 m.y.a.

During the last decade, methods for detecting DNA polymorphisms have proliferated at a bewildering pace. Today the investigator must choose among various types of genetic markers as well as between a variety of methods for discovering and screening polymorphisms. Polymorphisms useful for the analysis of population structure are found in both mitochondrial and nuclear genomes. Marker development may proceed along two routes: 1) discovery of species–specific markers, and 2) application of universal methods. Species-specific markers are based on sequence data from the target species or close relatives, whereas universal markers are based on the use of polymerase chain reaction (PCR) primers targeted to regions highly conserved across diverse taxa. Markers commonly employed include mitochondrial DNA polymorphisms, microsatellites, anonymous nuclear loci and known genes (both coding and noncoding regions). Methods for detecting polymorphisms range from technically simple (RFLP analysis) to more sophisticated mutation scanning methods. We review the application of these approaches to several key Antarctic species (the Patagonian toothfish Dissostichus eleginoides, the mackerel icefish Champsocephalus gunnari, and the squid Martialia hyadesi Rochebrune & Mabille, 1889) and present preliminary data on genetic polymorphisms in toothfish and icefish.

Community analyses of the macrobenthos living on the Weddell Sea shelf revealed a distinct horizontal patchiness. Within some systematic groups a specific faunistic classification could clearly be defined, e.g. for asterozoans and holothurians. Whilst for fish, only a general zoogeographical pattern was discernible, there were some recognisable relationships to different microhabitats. The extreme differences in the distribution of sponges observed seems to reflect their highly variable biological characteristics. Studies using underwater imaging methods for benthic research have provided strong evidence for the ecological significance of two factors. The first, iceberg scouring, leads to a variety of simultaneous stages of recolonization, which result in an increase in beta-diversity. As a consequence, it is unlikely that regionally a stage approaching a theoretical climax will ever be attained. Secondly, the structural diversity of living substrata provides the basis for an additional variety of epibiotic species. Only weak or non-detectable correlations have been found between benthic assemblages and physical parameters, such as water depth, sediment type, bathymetric features and the abundance of deposited phytodetritus. This indicates a benthic system which is relatively uncoupled from processes in the water column. The combination of stable environmental conditions and disturbances taking place over long periods of time, which are partly a special feature of Antarctica's glacial history, shaped the diversity and faunal composition of the macrobenthos. Consequently, neither Houston's “intermediate-disturbance-hypothesis” nor Sander's “stability-time-hypothesis” can be rejected for this part of the Antarctic ecosystem.

The terrestrial vegetation in Antarctica is restricted to small, very isolated, ice-free areas on the continent and on the islands adjacent to the Peninsula region. Lichens, a symbiosis between a fungus and an alga or cyanobacterium, form the most prominent component in the vegetation. They have the greatest number of species compared with other cryptogamic and unicellular organisms. They are able to colonize all types of solid substrata and have a low mineral nutrient demand. Their physiological vigour is derived from high freezing tolerance and the ability to be photosynthetically activated by water vapour uptake from snow at temperatures as low as c. -20°C. Long-term monitoring in their natural habitats demonstrates that lichens are photosynthetically active at suboptimal temperatures. Locally, however, they can benefit from meltwater on insolated rocks, but only for short periods in early summer. Although at this time they may be exposed to substrate temperatures of > 20°C and strong light they do not suffer from photostress. If, in winter, lichens are covered and kept dark by snow with temperatures close to 0°C, lichens enter a negative carbon balance. This effect may be relevant to lichens under conditions of global or regional warming.

Molecular analysis of phytoplankton population structure has lagged behind other groups and has usually been inferred from physiological data determined from relatively few clones. Nearly every physiological measurement has shown that no single clone of any phytoplankton species can be considered truly representative of that species. One important reason why studies of phytoplankton population structure are perhaps 20 or more years behind those of other organisms is because of the necessity to establish clonal cultures prior to genetic analysis and the inability to perform fine-scale sampling under most conditions. Isozyme analysis, performed for a few species, has revealed heterozygosity between populations. In addition, fingerprinting analyses, such as Random Amplified Polymorphic DNAs (RAPDs) or multi-locus probes, have shown that phytoplankton blooms are not mono-clonal, are highly diverse and isolates are related by geographic origin. In the Southern Ocean, only two studies have been made of the population structure of phytoplankton. The first, based on quantitative genetic analysis of morphometric features, suggests that there is sufficient genetic variation in populations of Thalassiosira tumida to allow speciation in terms of major shifts in morphology under conditions of continued directional selection. The second, using sequence data from the noncoding regions of the internal transcribed spacer region (ITS) in the ribosomal cistron as a molecular marker, shows that populations of Phaeocystis antarctica within continental water masses are homogenous with little evidence of population structure. Populations found within the Antarctic Circumpolar Current are genetically distinct from others, suggesting the currents also play an important role in determining population structure in phytoplankton populations.

The majority of species of the Antarctic coastal fish fauna is represented today by the perciform suborder Notothenioidei. The separation of basal lineages of notothenioids has been estimated to have occurred between 23 and 22 million years ago (m.y.a.), while a major diversification probably occurred 16–10 m.y.a. Cytogenetic approaches aim to study the genomic change that, at the cell level, accompanied the radiation of this group. The information available for 66 of 120–130 species makes possible the description of the main patterns of chromosome diversification within the suborder. Within some families (Channichthyidae, Artedidraconidae) the range of the chromosomal variability seems to be minimal whilst the high karyotypic diversity of Nototheniidae and Bathydraconidae is consistent with morphological and molecular data suggesting the paraphyletic nature of these two taxa. Molecular cytogenetics allows detailed chromosome characterization, including mapping of ribosomal genes and of telomeric sequences, thus providing information on processes of karyotypic rearrangement and direction of chromosomal change. Active process of genomic restructuring leads to intraspecific variability in several species, at different levels of chromosomal organization. The growing amount of information make it possible to use notothenioids as a model for testing hypotheses of evolutionary change in marine organisms, including chromosomal diversification.

A census of pelagic copepods identifies 346 species from 38 families (i.e. 16.7% of all copepods). Among them, 289 species are considered “cosmopolitan” and 57 “endemic”. Most of the species were described in the late 19th or early 20th century. However, due to a resurgence of interest in biological diversity, a large number of exhaustive revisions at various taxonomic levels have recently been initiated. One hundred and thirteen documents covering the census and distribution of copepod species are analysed. The distribution of species in the South Seas is given for 24 sectors chosen in relation to the geographical coordinates of the South Atlantic, South Pacific and Southern Indian oceans and their location with respect to the Antarctic Convergence or the continental shelf. The respective roles of Lagrangian displacement, of the hydrological features characterising the Southern Ocean and of the general ocean currents are emphasised to explain the scattered distributions of the species. Twenty-one species are catalogued as bipolar, 46 species as specifically sub-Antarctic and 222 cosmopolitan ones are probably from sub-tropical/temperate regions.

In investigating the extent of genetic diversity in Antarctic mosses the RAPD technique has proven especially useful in demonstrating that these mosses exhibit extensive genetic variation (levels being similar to those in temperate regions), and that within-colony variation is apparently caused not only by immigration and establishment of propagules from elsewhere, but also by mutagenesis. Dispersal of these mosses can also be followed using the RAPD technique; both short-distance dispersal by wind or water (depending on the moss species) and longer distance dispersal by wind across the ice cap has been demonstrated. Relationships with temperate mosses of the same species are being investigated to determine the origins of Antarctic populations. Genetic technology is also facilitating the taxonomic identification of moss specimens which cannot be identified morphologically due to phenotypic plasticity in Antarctica. Together, these genetic studies are starting to provide information of fundamental scientific importance for understanding the evolution, origins and dispersal mechanisms of Antarctic mosses, and their response to climate change.

Increasing interest in the ecological roles, conservation and biotechnological potential of Antarctic microbiota has focused attention on their biodiversity and evolutionary origins. Antarctic microbial ecosystems provide useful models for general questions in evolutionary ecology given the relative isolation of the South Polar Region, the severe biological constraints imposed by the polar environment, and the absence of higher plants and animals in some Antarctic habitats. Sealed environments such as Lake Vostok and the overlying East Antarctic ice sheet provide unique, natural culture collections for studying microorganisms that have been isolated from the global gene pool over timescales of evolutionary significance. Most Antarctic environments, however, continue to receive microbial propagules from outside the region, as indicated by spore trap data, the microflora found in Antarctic snow and ice, the colonising taxa at geothermal sites, and the high frequency of apparently cosmopolitan species in most habitats. Differences in environmental stability and selection pressure among environments are likely to influence the degree of adaptive radiation and microbial endemism. The latter seems greater in the Southern Ocean by comparison with non-marine ecosystems of Antarctica, although there is some evidence of endemic species in highly specialised niches on the continent such as in the endolithic habitat and saline lakes. Analytical techniques such as 16S rDNA sequencing and DNA–DNA hybridisation are beginning to provide new insights into the genetic affinities and biodiversity of Antarctic microbiota, and are leading to a more rigorous evaluation of microbial endemism.

The eight dominant culturable members of an Antarctic fellfield soil bacterial community were four Arthrobacter species, Sanguibacter suarezii, Aureobacterium testaceum, a Bacillus sp., and a Pseudomonas sp.. All of the isolates grew at 2°C, but two of the Arthrobacter spp. were psychrophilic, while the other six bacterial species were psychrotolerant. However, the fastest growing organisms at low temperatures were not the psychrophiles, and the psychrotolerant Bacillus sp. grew fastest at temperatures up to 25°C. When the growth temperature of cultures was altered, the phospholipid content of the two psychrophilic Arthrobacter spp. decreased, whereas the phospholipid contents of the psychrotolerant spp. either increased or did not change. Only one psychrophilic and one psychrotolerant Arthrobacter sp. modified its polar lipid head-group composition in response to a lowering of growth temperature. The change in Arthrobacter sp. CL2-1 was particularly marked and novel in that at low temperatures phosphatidylethanolamine was replaced completely by a phosphoglycolipid and phosphatidylserine, neither of which was present at higher growth temperatures. All eight isolates altered the fatty acyl compositions of their membrane lipids in a manner that was only partially dependent on taxonomic status. In Bacillus sp. C2-1 the changes were opposite to that predicted on the basis of membrane fluidity considerations. The isolates used different combinations of changes in fatty acid branching, unsaturation and chain length. There was no single strategy of thermal adaptation that was employed and the variety of strategies used did not follow phylogenetic boundaries.