ARTICLE IN PRESS Soil Biology & Biochemistry 38 (2006) 3003–3018 www.elsevier.com/locate/soilbio Diversity and distribution of Victoria Land biota Byron J. Adamsa,, Richard D. Bardgettb, Edward Ayresc, Diana H. Wallc, Jackie Aislabied, Stuart Bamforthe, Roberto Bargaglif, Craig Caryg, Paolo Cavacinih, Laurie Connelli, Peter Conveyj, Jack W. Fellk, Francesco Fratil, Ian D. Hoggm, Kevin K. Newshamj, Anthony O’Donnelln, Nicholas Russello, Rodney D. Seppeltp, Mark I. Stevensq a Microbiology and Molecular Biology Department, Evolutionary Ecology Laboratories, Brigham Young University, 775 WIDB, Provo, UT 84602, USA b Institute of Environmental and Natural Sciences, Soil and Ecosystem Ecology Laboratory, Lancaster University, Lancaster, LA14YQ, UK c Natural Resource Ecology Laboratory, Colorado State University, Fort Collins, CO 80523, USA d Landcare Research, Private Bag 3127, Hamilton, New Zealand e Department of Ecology, Evolution, and Organismal Biology, Tulane University, New Orleans, LA 70118, USA f Università degli studi di Siena, Dipartimento di Science Ambientali, Via Mattioli, 4, 53100 Siena, Italy g University of Delaware, College of Marine Studies, Lewes, DE 19958, USA h Dipartimento di Biologia Vegetale, Università ‘‘La Sapienza’’, Piazzale A. Moro, 5, 00185 Rome, Italy i School of Marine Sciences, University of Maine, 5735 Hitchner Hall, Orono, ME 04468, USA j British Antarctic Survey, Natural Environment Research Council, High Cross, Madingley Road, Cambridge, CB3 0ET, UK k RSMAS, University of Miami, 4600 Rickenbacker Causeway, Key Biscayne, FL 33149, USA l Dipartimento di Biologia Evolutiva, Università di Siena, via A. Moro 2, 53100 Siena, Italy m Center for Biodiversity and Ecology Research, Department of Biological Sciences, The University of Waikato, Private Bag 3105, Hamilton, New Zealand n Institute for Research on Environment and Sustainability, Devonshire Building, University of Newcastle, Newcastle upon Tyne, NE1 7RU, UK o Department of Agricultural Sciences, Imperial College at Wye, Ashford, Kent TN25 5AH, UK p Australian Antarctic Division, Channel Highway, Kingston, Tasmania 7050, Australia q Allan Wilson Centre for Molecular Ecology and Evolution, Massey University, Private Bag 11-222, Palmerston North, New Zealand Received 15 November 2005; received in revised form 11 April 2006; accepted 19 April 2006 Available online 16 May 2006 Abstract Understanding the relationship between soil biodiversity and ecosystem functioning is critical to predicting and monitoring the effects of ecosystem changes on important soil processes. However, most of Earth’s soils are too biologically diverse to identify each species present and determine their functional role in food webs. The soil ecosystems of Victoria Land (VL) Antarctica are functionally and biotically simple, and serve as in situ models for determining the relationship between biodiversity and ecosystem processes. For a few VL taxa (microarthropods, nematodes, algae, mosses and lichens), species diversity has been intensively assessed in highly localized habitats, but little is known of how community assemblages vary across broader spatial scales, or across latitudinal and environmental gradients. The composition of tardigrade, rotifer, protist, fungal and prokaryote communities is emerging. The latter groups are the least studied, but potentially the most diverse. Endemism is highest for microarthropods and nematodes, less so for tardigrades and rotifers, and apparently low for mosses, lichens, protists, fungi and prokaryotes. Much of what is known about VL diversity and distribution occurs in an evolutionary and ecological vacuum; links between taxa and functional role in ecosystems are poorly known and future studies must utilize phylogenetic information to infer patterns of community assembly, speciation, extinction, population processes and biogeography. However, a comprehensive compilation of all the species that participate in soil ecosystem processes, and their distribution across Corresponding author. Tel.: +1 801 422 3132; fax: +1 801 422 0519. E-mail address: bjadams@byu.edu (B.J. Adams). 0038-0717/$ - see front matter r 2006 Elsevier Ltd. All rights reserved. doi:10.1016/j.soilbio.2006.04.030 ARTICLE IN PRESS 3004 B.J. Adams et al. / Soil Biology & Biochemistry 38 (2006) 3003–3018 regional and landscape scales is immediately achievable in VL with the resources, tools, and expertise currently available. We suggest that the soil ecosystems of VL should play a major role in exploring the relationship between biodiversity and ecosystem functioning, and in monitoring the effects of environmental change on soil processes in real time and space. r 2006 Elsevier Ltd. All rights reserved. Keywords: Belowground; Biodiversity; Biogeography; Distribution ecology; Ecosystem services; Ecosystem functioning; Global change; Species diversity; Systematics; Taxonomy 1. Introduction Integrated studies of the relationship between soil biodiversity and ecosystem functioning are hampered by a dearth of fundamental taxonomic knowledge of soil organisms and their contribution to ecosystem functioning. It is apparent that, at least for macrofauna, functional diversity can influence important soil processes, such as decomposition (Heemsbergen et al., 2004; Roscher et al., 2004). However, the relationship between species richness and ecosystem functioning in soils remains unclear, in part due to the lack of taxonomic information, numerous species and limited studies that have investigated this relationship (see Mikola et al., 2002 for review). However, species richness should be of greater importance in lowdiversity ecosystems. A comprehensive compilation of all the species that participate in soil ecosystem processes, as well as how their distribution and abundance varies across variable spatial scales, is required (Wall, 2005). Such knowledge enables a better understanding and more realistic design of experiments that can determine how their interactions with the soil environment, and each other, facilitate ecosystem functioning. The vast majority of Earth’s soils are too biologically diverse to determine the functional role that each species plays in a food web. Although it is widely recognized that species, and not trophic groups, are the fundamental units of biology and ecology (Ghilselin, 1987; Bernard, 1992; Frost and Kluge, 1994; Wheeler, 1995; Adams, 1998; Brooks and McLennan, 2002), there have been a few major efforts made to recover all of the species in a given area. Two all taxa biodiversity inventories have been initiated in temperate North America (Great Smoky Mountains National Park; Mammoth Cave National Park). Others were conceived, but to date, the hyperdiversity of organisms smaller than 10 mm in each of these areas, and the lack of taxonomic expertise and infrastructure for dealing with these groups, poses an almost insurmountable task (Blackmore, 1996; Wheeler et al., 2004). Also, while recent initiatives have attempted to characterize a large portion of the soil biological community at a given site (e.g. Fitter et al., 2005), there are still no complete inventories for soil taxa. In species poor systems, such as Antarctica, richness is probably more important than other diverse systems, because there is less species redundancy, and a species inventory thus becomes feasible and a high priority. Victoria Land (VL), Antarctica is devoid of vascular land plants and hence feedback mechanisms between plants and soil biota cannot operate, as they do in other ecosystems (Wardle et al., 2004). Also, there are far fewer species in Antarctic soils than tropical and temperate ecosystems (Bargagli et al., 1997). Because of this, Antarctic soils potentially serve as excellent models for understanding fundamental properties of the relationship between soil biota and ecosystem functioning across multiple spatial scales (Barrett et al., 2004), and are one of few places on Earth where the taxonomic resolution necessary to finally address these questions is tractable. Here, we summarize what is presently known of the diversity and distribution of VL soil biota, and establish a baseline of taxonomic information to facilitate future efforts towards understanding their functional roles in soil ecosystem processes. For purposes of this assessment, we consider VL to be the part of Antarctica from the west side of the Ross Sea southward from 701300 S to 781000 S, and westward from the Ross Sea to the edge of the polar plateau (USGS, 2003). For ease of discussion we divide VL into two regions, northern Victoria Land (nVL) from about 701300 S to about 761S, encompassing Terra Nova Bay, Edmonson Point and Cape Hallett. We consider southern Victoria Land (sVL) to lie between 761S and about 781S including all of the McMurdo Dry Valleys and nearby coastal regions. The two areas correspond with a biogeographical split among some biota (i.e. Terra Nova Bay marks the southern limits of several taxa). In each of these regions, penguin rookeries, coastal areas and areas of moist and drier soils exist. Our coverage of biota includes bacteria, fungi, protozoa, rotifers, tardigrades, nematodes, mites, and springtails. Cyanobacteria and algae that occur in lakes and streams are included as the saturated sediments of lake margins and stream beds represent an important functional extension of the soil habitat (Treonis et al., 1999; Wall, 2006). Cryptoendoliths and cryoconites are other habitats for biodiversity, but their ecological linkages with soil biota are outside the scope of this paper, and are not considered in detail here. Our appraisal of the taxonomic diversity of soils in VL, which was explored only in the last hundred years, shows that identification and biogeographical distribution is limited by the same taxonomic restrictions shown for soil communities elsewhere. These include intensity of sampling effort across the various habitats within each region. ARTICLE IN PRESS B.J. Adams et al. / Soil Biology & Biochemistry 38 (2006) 3003–3018 3005 To date there have been no systematic surveys of bacterial diversity in the Dry Valleys. The soils of this region support a potentially diverse microbial community that for the most part is uncharacterized (Wynn-Williams, 1996; Cowan and Tow, 2004). Although there is relatively little detailed taxonomic information on the identities of soil microorganisms, there is a developing body of evidence for microbial activity in these soils (Burkins et al., 2001; Treonis et al., 2002; Parsons et al., 2004). Using the polymerase chain reaction, preliminary investigation has shown the presence of ammonia oxidising bacteria in soils of the Garwood Valley (A.G. O’Donnell, I.S. Waite, C.V. Lanyon, O. Owen, D.W. Hopkins, unpubl.) whilst molecular methods targeting the gene for methanogenesis specific methyl co-enzyme reductase (Hales et al., 1996) indicate the presence of methanogens between 20 and 50 m from lake margins (Hopkins et al., 2005). Similar genetic investigations of dry mineral soils from the Upper Miers Valley, Taylor Valley, and Cape Hallett (Luther Vale) show the microbial communities to be in low abundance yet highly diverse and heterogeneous (S.C. Cary and D. Cowan, unpubl.). Miers Valley soils appear dominated by Actinobacteria, a Proteobacteria, and Cyanobacteria until organically enriched when a completely different community (Bacilli and g proteobacteria) becomes dominant (L. Robson and S.C. Cary, unpubl.). These observations suggest VL dry soils harbor a more highly diverse microbial community than would be predicted by the extreme nature of the system. The environmental parameters that specifically drive this microbial diversity are currently unknown. lus and Pseudomonas. Coryneforms (Actinobacteria) were typically dominant in Ross Desert mineral soil whereas Pseudomonas and Bacillus were rare (Johnson et al., 1978). These studies led to the conclusion that bacteria from Antarctic soils may be cosmopolitan in distribution. 16S rDNA sequencing and phylogenetic analysis is providing new insights into the genetic affinities and diversity of Antarctic bacteria (Table 1). Consequently, using 16S rRNA sequencing some cultured bacteria have been assigned to known genera such as Arthrobacter and Bacillus whilst others, including Hymenobacter roseosalivarius (Hirsch et al., 1998) and Modestobacter multiseptatus have been recognized as belonging to novel genera (Mevs et al., 2000). Thermophilic bacteria have also been isolated from thermally heated soils near Mts Melbourne and Rittman in nVL (reviewed by Broady, 1993; Logan et al., 2000; Stevens and Hogg, 2006a). Notwithstanding the limitations of culture-based approaches to understanding community structure and diversity in soils (O’Donnell et al., 2005) comparisons of bacterial diversity and abundance between sites in these cold desert soils is compromised by the use of different media. Typically, the highest numbers of cultured bacteria are from relatively moist coastal soils, compared with the small bacterial communities of dry inland soils. However, using luminometry to estimate ATP levels, Cowan et al. (2002) concluded that the numbers of bacteria in cold desert mineral soil in Taylor Valley were higher than previously believed. The highest numbers of heterotrophic bacteria were cultured from nutrient-enriched ornithogenic soils of Cape Hallett (O’Brien et al., 2004) where bacteria tend to be most abundant at the surface and decline with depth. Similarly, higher numbers have been reported in samples at the interface of the soil and the ice-cemented permafrost (Vishniac, 1993). 2.1.1. Cultured bacterial diversity Molecular studies such as those outlined above that detect the presence of an amplifiable gene or fragment of a gene (DNA) do not necessarily establish that the organisms are intact and active. The cold and dry environment is a perfect setting to preserve DNA, in or out of the cell, indefinitely. More extensive molecular analyses (molecular probing or direct analysis of specific genetic transcripts), coupled with process related analyses (Griffiths et al., 2004; Malosso et al., 2004; Hopkins et al., 2005), are required to ascertain the viability of these bacteria. To definitively show that the organisms are active and capable of growth requires isolation and cultivation. To date most investigations of bacteria in soils of VL have focused on the abundance and diversity of cultivable bacteria in surface cold desert mineral soils of the Dry Valleys (reviewed by Vishniac, 1993). The bacteria described were aerobic heterotrophs that formed pigmented colonies on solid media. Anaerobic bacteria have not been widely reported. In early (pre 1990) studies the bacteria were assigned to known genera including Arthrobacter, Micrococcus, Bacil- 2.1.2. Cyanobacterial diversity In contrast to the eubacteria, the diversity of the cyanobacteria has been more extensively studied though we still lack detailed understanding of biogeographic distribution and relative abundance. Surveys of cyanobacteria date back to the beginning of the 20th Century and the expeditions of Shackleton (1907–1909; West and West, 1911) and Fritsch (‘Discovery Expedition’, 1912). Cyanobacteria are found in all types of aquatic habitats in VL and often dominate the microbial biomass of streams and lake sediments (Vincent, 1988). They are the most important mat-forming component in VL lakes and ponds. In particular, Leptolyngbya frigida (Fritsch) Anagnostidis and Komárek (Anagnostidis and Komárek, 1988) is dominant in all benthic mats of nVL and sVL (Seaburg et al., 1979; Wharton et al., 1983; Fumanti et al., 1995, 1997). This species is also frequently found in soils and as an epiphyte on mosses (Alfinito et al., 1998) revealing its high capability for adaptation. In VL soils, the cosmopolitan Nostoc commune Vaucher 1803 can, if supplied with a thin water film, develop to sizes 2. Biodiversity and distribution of Victoria Land biota 2.1. Prokaryotes ARTICLE IN PRESS 3006 B.J. Adams et al. / Soil Biology & Biochemistry 38 (2006) 3003–3018 Table 1 Bacteria cultured from soils of Victoria Land Division Genera or species Source GenBank accession # Cold desert mineral soils Actinobacterium Arthrobacter Brevibacterium antarcticum Modestobacter multiseptatus Pseudonocardia antarctica Cape Evans Harrows Peak Dry Valleys Asgard Range Dry Valleys AJ495808 O’Brien et al., 2004 AJ577724 Y18646 AJ576010 Firmicutes Paenibacillus Planococcus Cape Evans Cape Evans AJ495806 AJ496039 a-Proteobacteria Aquaspirilluma Azospirillum Brevundimonas Marble Point Wright Valley Cape Evans AF411851 and AF413109 AF411852 AJ495803 b-Proteobacteria Massila Cape Evans AJ496038 g-Proteobacteria Acinetobacter Pseudomonas Psychrobacter Stentrophomonas Cape Evans Marble Point Crater Circe Cape Evans Cape Evans AJ495807 AF411853-5 O’Brien et al. AJ495805 AJ495804 Hymenobacter roseosalivarius Chryseobacterium Sphingobacterium Dry Valleys Cape Evans Cape Evans Y18834 AJ495802 AJ496037 Planococcus Cape Hallett O’Brien et al. Bacillus fumarioli Bacillus thermoantarcticus Alicyclobacillus acidocaldarius Mt. Rittmann Mt. Melbourne Mt. Rittmann AJ250056 AJ493665 AJ493667 Bacteroidetes Ornithogenic soils Firmicutes Thermally heated soils Firmicutes All bacteria included in this table have been identified to genus level by 16S rRNA gene sequence analysis. a Two variants of the 16S RNA gene in this bacterium. visible to the naked eye (Cavacini, 2001). Although found as an epiphyte on nVL mosses (Alfinito et al., 1998) and in sVL soils (Holm-Hansen, 1964) these organisms are also widespread in aquatic environments in both regions (Howard-Williams et al., 1986; Fumanti et al., 1995). The genus Gloeocapsa is one of the few cryptoendolithic taxa with a high adaptation to extreme environmental conditions in rocks of the Dry Valleys (Friedmann and Ocampo, 1976; Friedmann et al., 1988). To date approximately 180 different cyanobacterial taxa have been recorded in VL (Fumanti and Cavacini, 2005) with aquatic environments showing the greatest diversity. 2.2. Algae Algae and cyanobacteria are present in almost all ice-free areas of VL and occur in soils, as epiphytes on mosses, in cyanobacterial mats and in plankton of lakes and ponds. It is also possible to find algae associated with rocks or living in the thin film of melted water in the snow patches. Early algal surveys in VL were carried out by Van Heurck (1909; Belgica expedition, 1897–1899), West & West (1911; E. Shackleton expedition, 1907–1909) and Fritsch (1912; Discovery Expedition, 1901–1904). Presently there are records of more than 300 algal taxa (Fumanti and Cavacini, 2005) belonging to the following divisions: Chlorophyta, Heterokontophyta (Bacillariophyceae, Xanthophyceae and Chrysophyceae), Dinophyta, Cryptophyta, and Euglenophyta, with the first two representing more than 97% of the total. As with the entire Antarctic continent, Bacillariophyceae (Diatoms) and Chlorophyta are the most widespread algae in VL. Diatoms are abundant in aquatic environments decreasing in number in terrestrial habitats, especially in soils of northern and southern VL (Seaburg et al., 1979; Cavacini, 2001). On the other hand, Xanthophyceae are almost absent from benthic cyanobacterial mats in both regions, but are an important component of the flora in soils of nVL (Cavacini, 2001). Chlorophyta are also important components of the mats in lakes and ponds of VL but tend to increase their relative importance in terrestrial environments and especially in soils, where they are the densest algal group (Holm-Hansen, 1964; Seaburg et al., 1979; Cavacini, 2001). Records from other algal groups (Dinophyta, Cryptophyta, and Euglenophyta) are mainly limited to freshwater communities of the Dry ARTICLE IN PRESS B.J. Adams et al. / Soil Biology & Biochemistry 38 (2006) 3003–3018 Valleys (Cathey et al., 1981; Alger et al., 1996; LaybournParry et al., 1997). The Dry Valleys have about 80 algal species, with 50 confirmed species of freshwater diatoms (Esposito et al., 2006). The VL algal diversity contains, based on present knowledge, a low number of endemics (Broady, 1996; Jones, 1996), probably due to the extensive use of European and North American literature for identifications (Sabbe et al., 2003). The most widespread species are, among the Heterokontophyta, the diatom Pinnularia borealis Ehrenberg 1843 which shows a uniform distribution in nVL from Cape Adare to Granite Harbour, mostly in association with sediments but also as an epiphyte on mosses (Broady, 1987). The cosmopolitan diatom Hantzschia amphioxys (Ehrenberg) Grunow in Cleve et Grunow 1880 tolerates a wide range of environmental conditions being recorded from high altitude sites (Mt. Feather, Barrett et al., 1997) to ponds located close to the sea in Cape Adare (Fritsch, 1912). Luticola muticopsis (Van Heurck) Mann in Round et al. 1990 previously described as endemic, has a homogeneous distribution in VL, but this needs further investigation as to its taxonomic stability. Kentrosphaera facciolae Borzi 1883 and Prasiola crispa (Lightfoot) Meneghini 1838 (Chlorophyta) are also key VL taxa. K. facciolae has a distribution limited to cyanobacterial mats and soils in nVL (Fumanti et al., 1997; Cavacini, 2001), whereas P. crispa is often recorded in southern and northern VL soils (West and West, 1911; Broady, 1985), as well as being widespread across the maritime and sub-Antarctic, and also known from the Arctic. Species of Xanthophyta show more restricted distributions. Xanthonema exile (Klebs) Silva 1979 is widespread in nVL soils (Cavacini, 2001) whereas Xanthonema bristolianum (Pascher) Silva 1979, Monodus coccomyxa Pascher 1939 and Heterococcus moniliformis Vischer 1937 are known only from Dry Valley lakes and soils (Seaburg et al., 1979). 2.3. Yeasts and filamentous fungi The yeasts and filamentous fungi encountered in VL soils are in most cases cosmopolitan, globally distributed species. In the majority of studies on these organisms, the fungi were isolated and cultured on agar media and identified by classical taxonomic methods, rather than by analysis of nucleic acid sequence data. The commonly isolated fungi fall within the zygomycetes and the anamorphic or teleomorphic ascomycetes and basidiomycetes. In nVL, the most frequently isolated filamentous soil fungi are species of Alternaria, Aspergillus, Cladosporium (typically C. cladosporioides (Fresen.) G.A. de Vries and C. herbarum (Pers.) Link), Geomyces (almost exclusively G. pannorum (Link) Sigler and J.W. Carmich., often reported as Chrysosporium pannorum), Phialophora (typically P. fastigiata (Lagerb. and Melin) Conant, often reported as Cadophora fastigiata), Phoma (often P. herbarum Westend.), Thelebolus microsporus (Berk. and Broome) Kimbr. 3007 and Mortierella antarctica Linnemann (Del Frate and Caretta, 1990; Zucconi et al., 1996; Fenice et al., 1997; Tosi et al., 2002, 2005). Chaetomium gracile Udagawa is frequently isolated from geothermally heated soil on Mount Melbourne in nVL (e.g. Broady et al., 1987; Zucconi et al., 1996). Filamentous fungi are more commonly isolated from soils in coastal regions, with many species recorded from bryophyte-dominated habitats around Terra Nova and Wood Bays by several Italian expeditions. Fewer species of yeasts than filamentous fungi have been isolated from nVL soils, but species of Cryptococcus (typically C. albidus (Saito) C.E. Skinner) and Rhodotorula (typically R. minuta (Saito) F.C. Harrison) are regularly isolated from soils in the northern region (Del Frate and Caretta, 1990; Tosi et al., 2002, 2005). In sVL, where there have been relatively few studies on filamentous soil fungi, species of Aspergillus (e.g. Aspergillus ustus (Bainier) Thom and Church) and Penicillium (e.g. Penicillium jensenii K.M. Zalessky) are commonly isolated from soil (Sugiyama et al., 1967; Baublis et al., 1991). Less frequently isolated filamentous fungi from southern VL soils are Phialophora spp., Cladosporium spp. (typically C. cladosporioides) and G. pannorum (Baublis et al., 1991; Onofri et al., 2000; Tosi et al., 2002). The diversity of filamentous fungi in sVL soils is apparently lower than that in the northern region (cf. Baublis et al., 1991; Tosi et al., 2005), but it is unclear whether filamentous fungal diversity is actually lower in the southern region, or the difference is owing to higher sampling effort in nVL. The diversity of yeasts, by contrast, is apparently higher in sVL soils (cf. Atlas et al., 1978; Tosi et al., 2005). The yeasts Aureobasidium pullulans (de Bary) G. Arnaud, Debaryomyces hansenii (Zopf) Lodder and Kreger, Leucosporidium scottii (Fell, Statzell, I.L. Hunter and Phaff) and species of Candida (often C. psychrophila (Goto, Sugiy. and Iizuka) S.A. Meyer and Yarrow), Cryptococcus (frequently C. albidus) and Rhodotorula (often R. rubra (Schimon) F.C. Harrison but also R. minuta) are commonly isolated from sVL soils (Tubaki, 1961; Sugiyama et al., 1967; Cameron et al., 1977; Atlas et al., 1978; Baublis et al., 1991; Vishniac, 1996; Connell et al., 2006). Yeasts are the only forms of soil fungi isolated from some soils in the Dry Valleys (Victoria and Wright Valleys) (Atlas et al., 1978; Vishniac, 1996). One species, Cryptococcus vishniacii Vishniac and Hempfling, appears to be indigenous to upland soils of the Dry Valleys (Vishniac, 1996). Animal pathogens in the basidiomycete yeast genus Trichosporon (T. cutaneum (Beurm., Gougerot and Vaucher) N. Ota and T. beigelii (Küchenm. and Rabenh.) Vuill.) have also been recorded from sVL soils (Tubaki, 1961; Baublis et al., 1991). The ecological role of most of the fungi listed above is in the primary decomposition of plant material. Most filamentous fungi utilize simple plant-derived carbon sources such as cellulose (Fenice et al., 1997), and yeasts such as Cryptococcus spp. are able to utilize simple sugars (Vishniac and Baharaeen, 1982), which may explain the ARTICLE IN PRESS 3008 B.J. Adams et al. / Soil Biology & Biochemistry 38 (2006) 3003–3018 greater fungal diversity in soil close to bryophyte-dominated areas in nVL. The observation of Baublis et al. (1991) that the highest abundance of fungi in Dry Valley soil was under moss also corroborates this view. Most of the fungi are incapable of decomposing lignin, which may reflect the absence of the polymer from uninhabited VL ecosystems. Several fungi, notably G. pannorum, are strongly keratinophilic (Mercantini et al., 1989), and frequently occur in ornithogenic soil (Onofri et al., 2000). A restricted number of fungi are found in consistent association with other organisms: notable examples are the springtail-capturing Arthrobotrys ferox Onofri and Tosi, the entomophthoralean fungus Conidiobolus antarcticus Tosi, Caretta and Humber, found in the southern region (Tosi et al., 2004a), and the nematode-trapping A. tortor Jarow. and Nematoctonus sp. in nVL (Onofri and Tosi, 1992; Tosi et al., 2004b). Fell et al. (2006), have performed the only molecularbased study of fungal diversity in VL soils. Analyzing sequences of large and small subunit rDNA extracted from Dry Valley soil they found Cryptococcus and Trichosporon spp. corroborating culture-based studies. These authors also noted the presence of fungal taxa different from those recorded by culture-based methods, such as nematodetrapping fungi in the genus Hohenbuehelia and the saprotrophs Phaeosphaeria sp., Coniochaeta ligniaria (Grev.) Massee and Cochliobolus heliconiae Alcorn. Many of the fungal species present in VL soils are cosmopolitan in nature, however some species and clades are unknown (Fell et al., 2006), which suggests the presence of an endemic mycoflora. 2.4. Lichens and mosses The first lichen collections from continental Antarctica were made at Cape Adare on H.J. Bull’s Antarctic expedition to nVL in January 1895, by C. Borchgrevinck. The first moss collections in VL were also made by Borchgrevinck, in 1899, during the Southern Cross Expedition (Gepp, 1902; Darbishire, 1910). The National Antarctic Discovery Expedition 1901–1904 provided the earliest published account of the flora of sVL (Cardot, 1907; Darbishire, 1910), listing seven mosses and 15 lichens collected from Granite Harbour, Mt. Terror, Discovery Winter Quarters, McMurdo Sound, and the Western Mountains (Taylor Valley region). Moss collections from Cape Royds, Hut Point, Cape Irizar, Cape Barne, high moraines at Cape Royds and the Strand Moraines in McMurdo Sound were made by the British Antarctic Nimrod Expedition 1907–1909 (Cardot, 1910). Lichens were collected by the British Antarctic Terra Nova Expedition 1910–1913, from Cape Adare, Cape Sastrugi and Evans Cove (Darbishire, 1923). Further botanical exploration in sVL was not undertaken until the International Geophysical Year of 1958–1959. Most Antarctic lichens known at this time, including those collected during the Byrd expeditions, were enumerated by Dodge (1973) (see also Castello and Nimis, 1995). Greene (1967) studied collections of mosses and discussed general taxonomic difficulties while using collections to produce distribution maps that concentrated on the Ross Sea area (from about 801S to 701S). Numerous opportunistic collections from the Ross Sea region have been made since then, principally by American, New Zealand, Australian, British, Japanese and Italian scientists, but there are few published reports on the flora or vegetation (Hale, 1987; Seppelt et al., 1992, 1995, 1996, 1998; Seppelt and Green, 1998; Søchting and Seppelt, 2003). The most southerly regional records for moss were published by Wise and Gressitt (1965) from the east side of the Massam Glacier, and on the east side of the Barrett Glacier. Broady and Weinstein (1998) reported cyanobacteria, fungi, algae, and two lichen species (Lecidea cancriformis and Carbonea vorticosa) from the La Gorce Mountains. The VL bryophyte and lichen flora is not rich, and the degree of endemism is much lower than formerly supposed. The number of species and endemic elements are more numerous in the maritime Antarctic and there appear to be no endemic genera. The bryophyte flora of nVL includes at least 11 species of moss: Sarconeurum glaciale Cardot and Bryhn, Bryoerythrophyllum recurvirostre (Hedw.) Chen, Hennediella heimii (Hedw.) R.H. Zander, Syntrichia princeps (De Not.) Mitt.; Campylopus pyriformis (Schultz) Brid. (restricted to specific geothermal sites) (Skotnicki et al., 2001); Ceratodon purpureus (Hedw.) Brid.; Anomobryum (Bryum) subrotundifolium (A. Jaeg.) J.R. Spence and H.P. Ramsay, Bryum pseudotriquetrum (Hedw.) P. Gärtner, B. Mey. and Scherb.; Pohlia nutans (Hedw.) Lindb.; Grimmia plagiopodia Hedw.; Schistidium (Grimmia) antarctici (Cardot) L.I. Savicz and Smirnova 1, and one hepatic Cephaloziella varians (Gottsche) Steph. at Mt. Melbourne fumaroles, Edmonson Point, Harrow Peaks, and other icefree areas (Bednarek-Ochyra et al., 2000; Broady et al., 1987; Lewis Smith, 1997). In recent years a number of moss species recorded in VL have undergone nomenclatural changes: Grimmia antarctici is now more appropriately placed in the genus Schistidium; Didymodon gelidus has been synonymized with Didymodon brachyphyllus (Ochyra and Zander, 2002); and Bryum algens Cardot was synonymized by Ochi (1979) in B. pseudotriquetrum. The genus Bryum exhibits in Antarctica a perplexing range of morphological variation (Ochi, 1979; Seppelt and Kanda, 1986) and considerably altered features occur in aquatic forms (Seppelt, 1983a,b). Kaspar et al. (1982) reported cf. B. algens from sediments in Lake Vanda; this material should be referred to B. pseudotriquetrum despite the assertion by Webby et al. (1996) that there are flavonoid chemotype differences between material determined as B. algens and other specimens referred to B. pseudotriquetrum. Bryum subrotundifolium was transferred to the genus Anomobryum by Spence and Ramsay (2002). In VL this predominantly coastal species is usually a yellowish green but in shaded localities may be entirely ARTICLE IN PRESS B.J. Adams et al. / Soil Biology & Biochemistry 38 (2006) 3003–3018 green. Bryum argenteum Hedw. has been reported from a number of localities. Longton (1973) used material collected from near McMurdo Station in his global interpopulational studies of the species but whether or not this material was really B. argenteum and not A. subrotundifolium remains equivocal. Recent collections made from the Cape Hallett area in December 2004, and initially considered to be morphologically very close to B. argenteum and not to A. subrotundifolium, have been shown by molecular studies to be identical to any other Ross Sea region collections of A. subrotundifolium (Stevens et al., in press). The number of lichen species in VL (o100 spp.) is much lower than the number of lichen species reported for Arctic regions (north America and Greenland—ca. 470 spp.) at about the same latitude (Thomson, 1997). Ice-free areas in VL show a great diversity of habitats, characterized by wide ranges of water and nutrient availabilities and substrates (different types of granites, schistose metamorphites, amphibolites, basalt and volcanic rocks). Many icefree areas and especially those at low altitude and close to the coast have suitable climatic, topographic and edaphic conditions for the development of around 12 bryophyte species and about 57 lichen taxa, 51 of which were identified to species level by Castello (2003). Macrolichens such as Usnea sphacelata R.Br., U. antarctica Du Rietz, Umbilicaria decussata (Vill.) Zahlbr. or U. aprina Nyl., and communities of weakly- or non-nitrophilous lichens (e.g., Pseudephebe minuscula (Nyl. ex Arnold) Brodo and D. Hawksw., Rhizocarpon superficiale (Schaerer) Malme and R. geographicum DC, and several species of Acarospora and Buellia) are relatively widespread in coastal ice-free areas. Sites with substrates influenced by seabirds are colonized by well-developed communities of nitrophilous lichen species (e.g., Caloplaca athallina Darb., C. citrina (Hoffm.) Th.Fr., Candelariella flava (C.W. Dodge and C.E. Baker) Castello and Nimis, Lecanora expectans Darb., Physcia caesia (Hoffm.) Fürnrohr, P. dubia (Hoffm.) Lettau, Rhizoplaca melanophthalma (Ram.) Leuck. and Poelt, Xanthoria elegans (Link.) Th.Fr., X. mawsonii C.W. Dodge). A rich lichen flora (by continental Antarctic standards) has been described at Cape Hallett (Murray, 1963), Birthday Ridge (Kappen, 1985), Harrow Peaks (Lewis Smith, 1997), Crater Cirque, Cape King, Vegetation Island, Prior Island (Castello, 2003), with a large number of circumpolar or wide-distributed species, some of which (e.g., Buellia subfrigida May. Inoue, Lacanora physciella (Darb.) Hertel or Lecidea andersonii Filson) are known from geographically very distant areas. During the 1990s Italian researchers collected about 400 lichen samples from 41 localities between Hallett Peninsula and Starr Nunatak (between 721–761S and 1621–1701E). The nVL lichen flora contains at least 23 genera (all of them widespread in both hemispheres), and 57 taxa. Among the 51 identified species, 10 were continental endemic, 26 were Antarctic–subantarctic in distribution, one Antarctic–South American, 16 species were bipolar, 3009 and seven cosmopolitan. Most taxa are crustose (47), seven are foliose and three fruticose (Castello, 2003). The collection has also been used for the revision of some critical taxa such as yellow Acarosporaceae (Castello and Nimis, 1994a), Candelariella (Castello and Nimis, 1994b), Usnea (Walker, 1985) and Sarcogyne (Seppelt et al., 1998) and to prepare a database of nVL lichens and a preliminary key for their identification (Castello and Nimis, 2000). Taxonomic uncertainties and the lack of comprehensive surveys make any rigorous comparison of lichen and moss diversity between nVL and sVL exceedingly difficult. Most moss and lichen species in nVL possess physiological attributes to allow their colonization and survival throughout VL coastal ice-free areas (Green et al., 2000; Kappen and Schroeter, 1997; Kappen and Valladares, 1999; Pannewitz et al., 2005; Schlensog and Schroeter, 2000; Schlensog et al., 2004). The lichen diversity (about 31 species) of Ross Island (e.g. Longton, 1973) and Granite Harbour (Seppelt et al., 1995, 1996) is comparable to that in coastal areas of nVL with similar substrates. Among mosses only S. princeps and P. nutans have not been reported from sVL. However, Didymon brachyphyllus (Sull.) R.H. Zander (formerly known in Antarctica as D. gelidus Card.) seems to occur only in sVL. The liverwort C. varians (exiflora) is known from Botany Bay, Granite Harbour (Seppelt and Green, 1998). The main difference between ice-free areas in sVL and those in nVL is the reduced variety of habitats and lower free water availability in the former region. Most ice-free areas in sVL are localized far from the coast, have low free water availability, lack concentrations of breeding sea birds (nitrogen and sea salts input), and are less diverse geologically. While in nVL communities of epi-, chasmo-, and hypolithic lichens predominate on the widespread igneous and metamorphic substrates that are generally of low porosity, the porous sedimentary rocks of the Beacon Supergroup found in the extreme desert conditions of the Dry Valleys support a suite of endo- and chasmolithic species. Lichens fill the interstitial spaces beneath the surface of the porous rock and grow between and around crystals. They lose their distinctive morphology and usually do not form reproductive structures (i.e., their taxonomic identification becomes exceedingly difficult). On granitic or metamorphic substrates in the Dry Valleys, the occurrence of chasmolithic and crustose taxa is, to a large extent, determined by moisture availability (in the form of cloud condensates and snow). In the Dry Valleys the normally epilithic lichen species (Acarospora gwynnii C.W. Dodge and E.D. Rudolph, Buellia frigida Darb., B. grisea C.W. Dodge and G.E. Baker, B. pallida C.W. Dodge and G.E. Baker, C. vorticosa (Flörke) Hertel (formerly Carbonea capsulata (C.W. Dodge and G.E. Baker) Hale), Lecanora fuscobrunnea C.W. Dodge and G.E. Baker, L. cancriformis C.W. Dodge and G.E. Baker, and Lecidella (Lecidea) siplei (C.W. Dodge and G.E. Baker) May. Inoue) are found primarily in protected niches beneath the rock surface occupying a cryptoendolithic ecological niche (Nienow and ARTICLE IN PRESS 3010 B.J. Adams et al. / Soil Biology & Biochemistry 38 (2006) 3003–3018 Friedmann, 1993). Although it was assumed that along VL environmental–climatic gradients, there are also gradients of lichen morphological forms (Hale, 1987; Hertel, 1988), geological features rather than climatic conditions determine the occurrence of cryptoendolithic communities in nVL. They appear to be widespread in areas where suitable porous sandstones occur (e.g. at Timber Peak), although they are conspicuously absent in sandstones at Vulcan Hills and Mt. Mackintosh. 2.5. Protozoa Protozoa in VL soils have thus far been studied by culture methods. The water-filled soil pore spaces generally have two domains: (1) the p8 mm ‘‘protozoan region’’ of small pores and pore spaces inhabited by flagellates and small amoebae; and (2) the 48 mm ‘‘protozoan-nematode’’ domain of larger pores and pore spaces inhabited by all groups of protozoa, nematodes, rotifers, and tardigrades (Bamforth, 1985). VL protozoa are essentially limited to flagellates, small amoebae and a few ciliates, and were found in 92% of samples collected from several Dry Valleys of sVL, where they were several orders of magnitude more abundant and diverse than nematodes. No endemic taxa have been found (Bamforth et al., 2005). Flagellates included Oikomonas termo Ehrenberg, Bodo saltans Ehrenberg, B. mutabilis Klebs, B. edax Kent and Heteromita globosa Stein. The small amoebae were of two types. The most abundant were Acanthamoeba and Echinamoeba, irregularly triangular amoebae with short subpseudopodia. The second group consisted of monopodal wormlike amoebae, the subcylindrical Hartmanella and Saccamoeba, and the lingulate Platyamoeba stenopodia Page. 2.6. Tardigrada Tardigrades (water bears) are widely distributed in soils, cryptogamic vegetation and freshwaters across Antarctica (Block, 1984; Convey, 2001), but few specific studies have been made on Antarctic (including VL) taxa. In VL, tardigrades are found primarily in moist or wet soils (Wall Freckman and Virginia, 1998). Identification by nonspecialists is difficult, and many taxonomic issues remain unresolved. Their presence in VL and elsewhere in the Ross Sea sector of Antarctica was noted by some of the earliest scientific expeditions (Richters, 1909; Murray, 1910), and during predominantly American terrestrial biological surveys during the 1960s (Armitage and House, 1962; Dougherty and Harris, 1963; Janetschek, 1967), but more thorough taxonomic evaluations of tardigrades from VL have been published over the two decades (Binda and Pilato, 1992, 1994, 2000; Pilato and Binda, 1990, 1997, 1999). Thus, reliable taxonomic knowledge is extremely limited, while distributional data are effectively restricted to the specific collection localities in which specialists have collected material or to one or two specific expeditions (e.g. Cathey et al., 1981). Other studies (e.g. Schwarz et al., 1993; Suren, 1990) often only give general reference to location (‘‘Victoria Land’’) or do not identify specimens beyond genus. Other than the critical taxonomic studies carried out in the region of Terra Nova Bay (nVL) most records would now appear to require taxonomic confirmation. These studies give the greatest recorded species diversity for this region (seven species; for comparison, approximately 18 tardigrade taxa have been reported across the continental Antarctic (Convey and McInnes, 2005)), but comparable data are lacking from elsewhere. One species (Acutuncus antarcticus Richters 1904) is currently recorded from both nVL and sVL. Species reported from nVL include Minibiotus furcatus Ehrenberg 1859, M. vinciguerrae Binda and Pilato 1992 (Binda and Pilato, 1992); Macrobiotus mottai Binda and Pilato 1994 (Binda and Pilato, 1994), Diphascon (Adropion) tricuspidatum Binda and Pilato 2000, A. antarcticus (Binda and Pilato, 2000), Ramajendas frigidus Pilato and Binda 1992 (Pilato and Binda, 1990), Diphascon (Diphascon) polare Pilato and Binda 1999, D. (D.) dastychi Pilato and Binda 1999 and D. (D.) victoriae Pilato and Binda 1999 (Pilato and Binda, 1999). Species reported from sVL (including some coastal areas that are close to, but not circumscribed within sVL as defined in the current paper—including Dailey Island, McMurdo Sound, Blue Lake, Terraced Lake, and Ross Island) include: A. antarcticus (Cathey et al., 1981); Macrobiotus sp., Hypsibius convergens Urbanowicz 1925, Hypsibius cfr mertoni simoizumii, Hypsibius (Diphascon) scoticus Murray 1905Diphascon (Adropoion) sp; Hypsibius (Diphascon) sp. (Janetschek, 1967); Macrobiotus arcticusA. antarcticus; Macrobiotus cfr polaris; Macrobiotus oberhäuseri Doyère 1840Ramazzottius oberhäuseri; Diphascon alpinum Murray 1906Diphascon (Diphascon) sp.; Diphascon (?) sp?Hebesuncus sp (Murray, 1910). From the McMurdo Dry Valleys area, Hypsibius alpinusDiphascon (Diphascon) sp.; Hypsibius arcticusA. antarcticus; Hypsibius oberhaeseriRamazzottius sp.; Macrobiotus polaris (Dougherty and Harris, 1963) have been recovered. Porazinska et al. (2004) found tardigrades (A. antarcticus and Hypsibius spp.) in cryoconite holes of glaciers in Taylor Valley, always in the presence of rotifers. It should be emphasized that most of these records have not been subjected to recent critical taxonomic re-evaluation. 2.7. Rotifera Murray (1910) provided one of the earliest reports of rotifers in VL, observing 15 species (four Monogononta and 11 Bdelloidea) in lakes of McMurdo Sound area, sVL. Further sampling of the lakes in this area added only a few additional species (Armitage and House, 1962; Dougherty and Harris, 1963; Spurr, 1975; Laybourn-Parry et al., 1997). Porazinska et al. (2004) reported two rotifer species (Philodina gregaria Murray 1910 and Cephalodella catellina Muller 1786) from cryoconite holes located on five glaciers ARTICLE IN PRESS B.J. Adams et al. / Soil Biology & Biochemistry 38 (2006) 3003–3018 (Commonwealth, Canada, Howard, Hughes and Taylor) in sVL. However, despite the apparent similarities between cryoconite sediment and soil, the biotic community consisted of species typical of lake and stream sediment (Porazinska et al., 2004), thus the occurrence of these rotifer species in Antarctic soils remains uncertain. In a study that identified terrestrial rotifers to species in VL, three species (Philodina sp., Habrotrocha sp. and Epiphanes sp.) were found in moss-dominated moist soil near Canada Glacier, sVL (Schwarz et al., 1993). Despite the paucity of data on rotifer species occurrence, it is evident that as a taxon rotifers are widely distributed across VL soils, having been observed in southern (e.g. Wall Freckman and Virginia, 1998) and northern regions (Adams and Wall, unpubl.), as well as on nearby islands (Sinclair and Sjursen, 2001; Porazinska and Wall, 2002). Clearly we currently have very limited knowledge of the diversity of rotifers in VL. Several studies have reported total numbers of rotifers (albeit typically not identified to species) across a range of habitats and environmental gradients. However, since all published studies are limited to sVL it is impossible to determine broad scale distributional patterns. Rotifers, like most terrestrial invertebrates in Antarctica, are patchily distributed. For instance, they were found in only 4–14% of soil samples collected from the Dry Valleys, sVL (Wall Freckman and Virginia, 1998) and, although they were consistently found in penguin rookeries on Ross Island, near sVL, there was much spatial and temporal variation in abundance within each rookery (Porazinska and Wall, 2002). Soil moisture appears to be an important driver of rotifer abundance; Schwarz et al. (1993) reported a threefold increase in their abundance after glacial melt water had moistened soil and Treonis et al. (1999) observed greater populations of rotifers in stream sediments than in soils 8–32 m from the stream. In contrast, Sinclair and Sjursen (2001) found no relationship between rotifer abundance and soil moisture, but a positive correlation with soil organic matter was observed. These differences may be due to differences in habitat between coastal nVL and the Dry Valleys of sVL. In cryoconite holes rotifer abundance decreased with elevation, and variation in rotifer abundance was explained by differences in pH, nutrient availability and hole diameter (Porazinska et al., 2004). At a larger scale (10–50 km), Courtright et al. (2001) demonstrated variation in rotifer abundance along three valleys in sVL, generally with greater abundances further up the valley although it is not clear what caused this pattern. 2.8. Nematoda The British Discovery expedition of 1901–1903 yielded the first nematode specimens from VL, collected near Discovery Bay, sVL. Steiner (1916) later described the specimens as Dorylaimus antarcticus (syn. Eudorylaimus antarcticus Yeates, 1970) but over 50 years passed before taxonomic work on the nematodes of VL was resumed (Yeates, 1970; Timm, 1971). 3011 Nematodes are the most commonly encountered and abundant metazoans of VL soils (Wall Freckman et al., 1997; Wall Freckman and Virginia, 1998). However, identification by non-specialists is difficult, and the taxonomic status of many VL species is currently unresolved. The most geographically ubiquitous and taxonomically stable of the VL soil nematodes is Scottnema lindsayae Timm 1971. S. lindsayae is present throughout VL where it tolerates a wide range of soil moisture and geochemical conditions. Species of the genus Plectus have been reported from north and south VL; P. antarcticus de Man 1904 Timm 1971, P. frigophilus Kirjanova 1958, and P. accuminatus Bastian 1865. All three purportedly occur in coastal nVL (Terra Nova Bay; Vinciguerra, 1994) whereas only P. antarcticus and P. accuminatus have been reported from sVL where they are common in wet soils near streams and lakes. However, it has been suggested that P. antarcticus occurs only in maritime Antarctica, and that specimens identified as P. antarcticus in VL are P. murrayi Yeats 1970 (Andrássy, 1998; see also Maslen and Convey, this volume). E. antarcticus Steiner 1916, E. shirasei Kito, Shishida and Ohyama 1996, and E. glacialis Andrássy 1998 have been reported throughout VL where they occur in moist soils and stream and lake sediments. However, the taxonomic identity of E. antarcticus reported from ecological studies done in the Dry Valleys of sVL has been questioned, with the suggestion that they are more likely E. glacialis or E. shirasei (Andrássy, 1998). Geomonhystera antarcticola Andrássy 1998 has been recovered from moist soils in sVL, but it is not yet clear whether these populations represent one or more species. They are very rare and patchily distributed in the Dry Valleys of sVL, and occur in greater abundance in the wetter, more productive coastal nVL soils of Edmonson Point and Cape Hallett. Panagrolaimus davidi Timm 1971 has been recorded primarily from coastal regions of VL, and is both common and abundant in moist, productive environments. It is only rarely encountered in Dry Valley soils of sVL. In the Dry Valley soils of sVL, Plectus, Eudorylaimus and Scottnema have been shown to have distributions that are sensitive to available moisture, pH, EC (electrical conductivity), and inorganic C (Freckman and Virginia, 1997; Powers et al., 1998; Treonis et al., 1999; Virginia and Wall, 1999; Courtright et al., 2001; Porazinska and Wall, 2002). The definitive taxonomic treatise on the diversity of VL nematofauna remains the treatment of Andrássy (1998). However, constraints on the condition, number, and geographic representation of the specimens examined by Andrássy (1998) indicate the need for careful taxonomic studies that encompass a larger number of specimens sampled across a broader range of environmental gradients, particularly for Plectus, Geomonhystera and Eudorylaimus. 2.9. Arthropoda Springtails (Collembola), and mites (Acari), are the largest endemic terrestrial invertebrates, indeed the largest ARTICLE IN PRESS 3012 B.J. Adams et al. / Soil Biology & Biochemistry 38 (2006) 3003–3018 year round terrestrial animals, found in VL and the Queen Maud Mountains (Wise, 1967; Stevens et al., 2002a). They have received attention from a number of researchers owing to their relatively conspicuous nature and presence in locations of historical importance. For example, some of the first springtails collected in VL were during the British Antarctic Expedition, 1898–1900, and again during the British National Antarctic Expedition, 1901–1904 (see Wise, 1967; Stevens et al., 2006a). Accordingly, their diversity is relatively well known, and their distribution is known to be limited to areas of high soil moisture and/or access to water, such as streams, or snow meltwater (see Kennedy, 1993; Hogg and Stevens, 2002). They are found throughout nVL and sVL (including the Dry Valleys), and as far south as the Queen Maud Mountains (see also Sinclair and Stevens, 2006). Their taxonomic diversity is limited in VL (seven species of springtail and ten species of mites), although three species of springtail (all endemic) and four species of mites (two endemic) are found in the less studied vicinity of the Queen Maud Mountains. Distinct biogeographic breaks occur in the distribution of arthropods in VL, and indeed the Ross Dependency. The Drygalski ice tongue corresponds to a major biogeographic break separating northern from southern VL biota. Northern VL contains the greatest species diversity for both springtails and mites. Of the ten mite species in nVL, six are endemic (three are cosmopolitan) and three are shared with sVL, where only one of the four sVL species is endemic (Coccorhagidia keithi). The two main species of springtails found in nVL are Desoria klovstadi Carpenter 1902 and Gressittacantha terranova Wise 1967, both belonging to the Isotomidae. The distribution of D. klovstadi covers the northern part of the VL, from Cape Adare to the northern margins of the Mariner Glacier (Wise, 1967, 1971; Frati et al., 1997; Stevens et al., 2006a). The presence of D. klovstadi ends abruptly at the Mariner Glacier, and south of this glacier, it is replaced by G. terranova as the most abundant collembolan. The distribution of G. terranova is mainly restricted to the area from the Mariner Glacier to the David Glacier, although a partial overlap with D. klovstadi has been reported with few specimens collected at Crater Cirque and Football Saddle (Frati et al., 1997). A detailed survey using genetic markers (Fanciulli et al., 2001) shows the populations of G. terranova are arranged in three genetically distinct groups, each one limited by major glacier systems (the Aviator and the Campbell Glaciers) and with very little gene flow among them. A third collembolan relatively common in nVL (but not as common as D. klovstadi or G. terranova) is the neanurid Friesea grisea Schäffer 1891. Its distribution is scattered, and records are available from Cape Hallett in the north (Wise, 1967, 1971), to the Tinker Glacier in the south (Frati et al., 1997), but it is also known from outside the region (see Stevens and Hogg, 2006a). The fourth collembolan species inhabiting nVL is the isotomid Cryptopygus cisantarcticus Wise 1967. This species has been recorded from the Cape Hallett area (Wise and Shoup, 1967), Crater Cirque and Cape Phillips (Wise, 1967, 1971), but in terms of abundance it is always outnumbered by the sympatric D. klovstadi. In sVL, the springtail Gomphiocephalus hodgsoni Carpenter 1908 and the mite Stereotydeus mollis Womersley and Strandtmann, 1963 are by far the most widely distributed species. Both species show highest levels of genetic diversity in the McMurdo Dry Valleys (e.g. Stevens and Hogg, 2003; Stevens and Hogg, 2006). Two other species of springtail (Neocryptopygus nivicolus Salmon, 1965, Antarcticinella monoculata Salmon, 1965) and three other species of mite (C. keithi Strandtmann, 1967, Tydeus setsukoae Strandtmann, 1967, Nanorchestes antarcticus Strandtmann, 1967) have more restricted distributions, and in some cases are known only from one or two sites (e.g. Strandtmann, 1967; Wise, 1967, 1971). This pattern is repeated further south in the Queen Maud Mountains. Of the mites, three species are common across nVL and sVL, and two of these are also found further south in the Queen Maud Mountains. No springtails are common across these three regions (see Table 1, Sinclair and Stevens, 2006). 3. Discussion To date, biotic surveys of VL have focused on highly localized populations. Sampling effort has been greatest for the microarthropods, but even this has been targeted at relatively few locations in VL. The proportion of known to unknown taxa found in studies of other groups appears to decrease in relation to sampling effort and body size. Microeukaryotes and prokaryotes are two of the most understudied groups, yet are likely the most diverse. A similar trend is seen in respect to endemism. Species endemic to VL are rare; endemism is highest for microarthropods and nematodes, less so for tardigrades and rotifers, and (based on present data) very low for mosses, lichens, microeukaryotes and prokaryotes. The presence of a wide range of taxa in VL terrestrial ecosystems indicates that soil habitats are diverse in the region and vary spatially across broad and often steep environmental gradients. Several VL taxa appear to have localized distributions associated with highly specific soil properties. A general observation is that diversity appears to vary from low to high across a latitudinal gradient from sVL to nVL, and this is broadly associated with increased ecosystem productivity (Barrett et al., 2006). Another trend indicates an increase in diversity with available soil moisture, soil carbon and moderate salinities. For example, microarthropods and nematodes seem to follow these patterns of abundance and diversity across all three gradients. However, such generalizations are yet to be examined rigorously in functional or comparative community contexts (e.g. latitudinal differences in biodiversity are not observed among penguin rookery soils). Regardless of sampling intensity and taxonomic work, links between taxa and their functional contribution to ecosystems have not been adequately explored for the ARTICLE IN PRESS B.J. Adams et al. / Soil Biology & Biochemistry 38 (2006) 3003–3018 majority of VL taxa. VL soil food webs are likely among the simplest on the planet, yet the existence or importance of trophic relationships among the majority of taxa are still unknown, as are their net contribution to ecosystem functioning (see Hogg et al., this volume). Novel trophic relationships for metazoans are still emerging, and at present most of these relationships are only inferred or predicted based on similarities with better characterized, but non-indigenous taxonomic groups. In particular, trophic relationships between microscopic eukaryotes and other VL soil biota are very poorly known, yet could be major contributors to key soil processes. Future work will require careful observation of these taxa in situ (where possible), and will take advantage of approaches that trace resource pulses through food webs. Additionally, species’ contributions to ecosystem functioning vary spatially and temporally, and the distinction between functional (active) and non-functioning (inactive) taxa must be made. For example, numerous taxa may persist in a cryptobiotic state without contributing to ecosystem processes during a given environmental extreme (e.g., of altered temperature, moisture or salinity). Approaches that fail to distinguish active from inactive species artificially inflate the number of taxa involved in ecosystem functioning at any one time, and mask the role played by what may be more vulnerable species (including endemics), which are interpreted as being redundant. Existing studies of taxonomy and distribution serve largely to illustrate the gaps remaining in contemporary knowledge, rather than allowing biogeographical analyses and patterns to be used with any level of confidence. Furthermore, most of what is known of the biodiversity and distribution of VL biota exists in an evolutionary and ecological vacuum. Surveys and inventories to date have been taxon specific, and there have been no coordinated, integrated all-taxon inventories of these species as they exist in their respective communities. Taxon specific surveys are often devoid of any phylogenetic context (but see Stevens et al., 2006b), which restricts what can be logically inferred about their historical biogeography or how they came to be distributed and structured into their present communities. Syntheses of multi-species phylogeographies with geophysical variables will allow for more precise tests of what drives patterns of diversification and distribution of VL biota (for example, Lapointe and Rissler, 2005; Stevens et al., 2006b). There are clear community differences in some groups (particularly the arthropods) between nVL and sVL, although whether these can simply be related to latitudinal (proxy for environmental) gradients or are a more fundamental indicator of historical processes through vicariance remains an area of active research (e.g. Stevens and Hogg, 2003; Stevens et al., 2006b; Stevens and Hogg, 2006b). As yet, there remains a virtually complete absence of detailed autecological studies of specific taxa across all functional groups, with assessments of the existence and importance of ecological function largely being little more 3013 than inference, based on knowledge of related taxa or comparable communities elsewhere. Sampling effort has varied widely across groups, currently being greatest (but still inadequate) for microarthropods, mosses and lichens. Likewise, molecular taxonomic approaches have so far focused mostly on limited representatives of the microarthropods, and there is a clear and urgent need to apply these more widely, in concert with classical taxonomic and autecological approaches, across the various higher taxa. Such studies are an urgent requirement, not least, so as to provide the requisite baseline against which to monitor the consequences of regional and global change on ecosystems within VL. They will also allow accurate assessment in the virtually unaddressed field of cryptic speciation. Amongst the fauna, the ubiquity and abundance of protozoa (predominantly flagellates and small amoebae), compared to metazoans emphasizes their importance in food webs and nutrient cycling in this most extreme ecosystem on Earth. However, the metazoans have received considerably more attention, both in VL and more widely across the Antarctic. To exist in these extreme environments, fauna, algae, mosses and lichens have survival strategies that allow them to persist across a wide range of habitats, alone and as communities. Across the Dry Valleys, for example, well-developed communities of many invertebrate taxa can be found close to soils in which no invertebrates exist. These patterns also occur at broader scales, where 40% of the soils are without nematodes, and some soils harbor no fauna at all. Our present knowledge of the diversity and abundance of biota in VL is extremely limited, and in need of coordinated, rigorous sampling. Based on trends of previous efforts in temperate ecosystems, campaigns may (a) reveal new species, (b) aid in resolving taxonomic problems and (c) extend the known ranges of species further south. Even soil algae, which have the highest recorded diversity (about 300 taxa), have yet to be examined in many remote sites. The application of molecular phylogenetic techniques will doubtless considerably advance knowledge of faunal, algal, lichen and microbial phylogeny and cryptic speciation. Comprehensive biogeographical research will allow the recognition of hot-spots of biodiversity. Even where there is much known about the distribution on continental and polar scales, such as with lichens and mosses, there is a need for adding to the knowledge of the factors limiting distribution. The lichen and moss flora of VL is characterized by the occurrence of many circum-Antarctic species and, in spite of a high incidence of endemic lichens in the Antarctic as a whole (about 50%, Øvstedal and Lewis Smith, 2001), it also shows clear affinities with the Arctic flora (about 27% of bipolar elements for the lichens). However, these data are not definitive because of the relatively poor knowledge of species distribution and the taxonomic uncertainty of many taxa. Without the identification to species levels of all taxa and critical revision of many widespread genera (which are ARTICLE IN PRESS 3014 B.J. Adams et al. / Soil Biology & Biochemistry 38 (2006) 3003–3018 very polymorphic, and often have closely related counterparts in the Northern Hemisphere), it is impossible to compare the lichen and moss diversity and distribution in nVL and sVL ice-free areas. On the basis of current data, there seems to be very little systematic difference in lichen or moss diversity between nVL and sVL, except for the Dry Valleys region, which has peculiar geological and climatic features. A combination of classical and modern research techniques, including support for the preparation of specific monographs and reliable and user-friendly identification keys, the proper deposition of descriptions, and the curation and accessibility of type collections, all in close integration with molecular approaches (both phylogenetic and population genetic), are required and should attract and involve wider interaction across appropriate specialists at a global scale. Molecular tools will facilitate efforts to explore the diversity and function of these groups, but these tools need to be applied to all VL taxonomic groups in order to resolve nomenclatural problems, aid identification, reveal cryptic species, and provide information about population processes, such as effective size, structure, dispersal and gene flow. These data also are amenable to statistical and comparative phylogeographic studies, which can reveal the concerted response of species and communities to the historical environmental changes that have shaped their present distributions. In the near future, technical advances will allow expression arrays of numerous taxa to be established for some of the gene products involved in fundamental soil ecosystem processes, such as nitrogen mineralization, catabolism of carbon substrates, and sulfur reduction. With replication and rigorous statistical analysis, comparisons among species, and not just different soil communities, will become possible allowing assessment of the relative contributions of not just functional groups, but individual species, to ecosystem processes. Understanding the relationship between soil biodiversity and ecosystem functioning remains a huge obstacle to predicting the global effects of environmental change. Integrated studies of the relationship between soil biodiversity and ecosystem functioning generally requires a comprehensive and integrated compilation of the species that participate in soil ecosystem processes, their distribution, and limits to distribution, across regional and landscape scales, something which is currently achievable in VL with the resources, tools, and expertise currently available. Because VL is a microcosm of more diverse and complex temperate and tropical ecosystems, the knowledge gained would be of great benefit for biodiversity and ecosystem dynamics (Wall 2004, 2005). Additionally, given the sensitivity of polar ecosystems to environmental change (Freckman and Virginia, 1997; Convey, 2003; Wall, 2005, 2006), a comprehensive baseline knowledge of VL biota and how it is distributed will allow for the monitoring of environmental changes to VL soil ecosystems in real time and space. 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