Southern Crustacean Relicts

Eophreatoicus from Kakadu in the Northern Territory of Australia. Image from here.

This week's highlight taxon is the Phreatoicidea, a suborder of isopods restricted to freshwater habitats in ex-Gondwanan continents. This is not a particularly large group - only about a hundred species have been described, though it is estimated that at least that number again remain undescribed. A reasonably high proportion of the species are known from subterranean habitats*, including the first species to be described, Phreatoicus typicus. Knott (1986) listed eleven subterranean species, which at the time was about a quarter of the total known diversity (over half the known species have been described since then). The diversity of the suborder is also heavily centred in Australasia - 94 species have been described from there, in contrast to four species from South Africa and only two from India (Wilson, 2008), but again this is probably heavily biased by the fact that almost all taxonomic work on this group has been conducted in Australia. For instance, Knott (1986) refers to possible undescribed species from India - these species seemingly still have not appeared in print twelve years later. The supposed Gondwanan distribution of phreatoicids also makes their apparent absence from South America very interesting, but how confident can we be that they are truly absent from that continent?

*I should make it clear that "subterranean" does not necessarily mean "cave-dwelling". Caves actually only make up a small proportion of the subterranean habitat, and only one cave-dwelling phreatoicid species is known (Knott, 1986). The majority of subterreanean species are sediment-dwelling forms whose habitats can extend right down into groundwater aquifers. Phreatoicus typicus, for instance, was originally described from a well near Christchurch in New Zealand, into which it would have emerged from the surrounding bedrock.

Most people imagine isopods as dorsoventrally flattened animals, like their most familiar representatives the woodlice and slaters. Phreatoicids, however, represent an exception to this rule, being fairly high-vaulted, narrow animals. Stygobiotic forms tend to be more elongated. Phylogenetically, phreatoicids are one of the most basal groups of isopods, and have one of the earliest fossil records. The Palaeozoic phreatoicids (or, technically, stem-phreatoicids) Palaeophreatoicidae are known from marine sediments, but since the Triassic all known representatives have been freshwater. Phreatoicids are detritivores feeding primarily on decaying vegetation or on the micro-organisms associated with the former, but may occassionally be carnivorous (Wilson, 2008). Phreatoicids are a significant part of the pholeteros - the specific faunal assemblage of organisms associated with the burrows of larger animals such as freshwater crayfish.


Pilbarophreatoicus platyarthricus, a potentially subterranean form from the Pilbara in Western Australia. Pilbarophreatoicus was described from an intermittent stream (i.e. one that dries up outside the rainy season), but shows features usually associated with subterranean habitats such as blindness and elongated body form. Like many subterranean phreatoicids in arid regions, it probably emerges from the ground when standing water is available and retreats back into the groundwater during the dry season. Figure from Knott & Halse (1999). shale bar = 1 mm.

Taxonomically, the Phreatoicidea have been a difficult group. For many years the classificatory sytem used was that established by G. E. Nicholls in the early 1940s, which divided phreatoicids between two families, each divided into a number of subfamilies. Unfortunately, the features used to separate these taxa have been shown to be largely artificial, and a high degree of variation can occur between closely-related species or even within examples of the one species. Wilson & Keable (2002)revised the classification somewhat through phylogenetic analysis, recognising three families and abandoning Nicholls' subfamilies. The suborder as a whole seems to be characterised by fairly slow morphological evolution. Gouws et al. (2004) showed that at least one supposed species, the South African Mesamphisopus capensis, is divisible on genetic and morphometric grounds into a number of potential cryptic species.

Like many freshwater and subterranean organisms, many phreatoicids have very restricted distributions and are placed at significant risk of human activities. Genetic studies show that each separate aquifer may have its own isolated population (Wilson, 2008). Indeed, a number of species are believed to have already gone extinct, due to factors such as increasing groundwater salinity as a result of deforestation (Knott, 1986) or alteration and exhaustion of water supplies and aquifers (Wilson, 2008). Unfortunately, the lack of taxonomic resolution within the group, as well as the difficulty of surveying the habitats of subterranean species in particular, make it very difficult to assess the risk to individual species.

REFERENCES

Gouws, G., B. A. Stewart & S. R. Daniels. 2004. Cryptic species within the freshwater isopod Mesamphisopus capensis (Phreatoicidea: Amphisopodidae) in the Western Cape, South Africa: allozyme and 12S rRNA sequence data and morphometric evidence. Biological Journal of the Linnean Society 81: 235-253.

Knott, B. 1986. Isopoda: Phreatoicidea. In Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) (L. Botosaneanu, ed.) pp. 486-492. E. J. Brill / Dr. W. Backhuys: Leiden.

Knott, B., & S. A. Halse. 1999. Pilbarophreatoicus platyarthricus n.gen., n.sp. (Isopoda: Phreatoicidea: Amphisopodidae) from the Pilbara Region of Western Australia. Records of the Australian Museum 51: 33-42.

Wilson, G. D. F. 2008. Global diversity of Isopod crustaceans (Crustacea; Isopoda) in freshwater. Hydrobiologia 595: 231–240.

Wilson, G. D. F., & S. J. Keable. 2002. New Phreatoicidea (Crustacea: Isopoda) from Grampians National Park, with revisions of Synamphisopus and Phreatoicopsis. Memoirs of the Museum of Victoria 59 (2): 457-529.