Volume 30(2), 2003

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Subterranean aquatic Crustacea Victoria Cave, Yorkshire, UK Stone money in Micronesia Planinska jama, Slovenia Cover photo by Phil Chapman: Androniscus dentiger Verhoeff, 1908 (Crustacea: Isopoda), commonly known as the Rosy Woodlouse. This animal is a troglophile and is one of the commonest woodlice to be found in British caves. It is a detritivore, which means that it can exist on food from diverse sources. No studies of this species have been carried out to determine its exact ecological status in British caves. However, extensive work by Gentile and Sbordoni in Italian caves (e.g. Evolution, Vol 52, 432-442, 1998) has demonstrated clearly that individual populations of this species, in separate caves, are very distinct genetically, and do not exchange genes with other cave populations or with surface populations. Thus, each population is effectively troglobitic and effectively a separately evolving taxon. Given the findings of this study, it seems very likely that many populations of cave animals (and perhaps especially terrestrial ones) will be found to be independently evolving entities. In British caves and groundwater it is highly likely that many genetically distinct populations exist. It is equally likely that British populations are distinct genetically from those in mainland Europe. This topic is discussed in a Paper by Proudlove et al in this Issue.

Editorial

(Gunn, J. & D. Lowe (eds))
Editorial
Vol 30 (2) pp 51 - 52

This Issue contains a Paper that is the first of what we hope will be several on related topics, so we have asked the lead author, Graham Proudlove, to write a Guest Editorial explaining the background and also presenting some potentially controversial thoughts on the future direction for biospeleology in Great Britain.
GUEST EDITORIAL (Proudlove, G.)
This Issue of Cave and Karst Science carries a Paper on some aspects of the biology of cave and groundwater animals. This is not unique. In recent years there have been numerous biological papers in Cave and Karst Science, but this one is notable. It is based on data collected during the heyday of cave biology in Britain and Ireland and is the first of a number using a remarkable and very valuable dataset. The editors asked me to write a Guest Editorial on subterranean biology to introduce this Paper. I will say a few things about the history of subterranean biology in the British Isles, about our current level of knowledge, and about how we progress from here. Some of what I say may be controversial. Thanks to Dave Lowe and John Gunn for giving up this space.
The study of subterranean biology in Britain and Ireland has a long and distinguished history. In 1938 a small group of cavers started to collect animals from caves, the main impetus being provided by E. A. Glennie and Mary Hazelton. It was Hazelton who took on two vital tasks: getting the animals accurately and authoritatively identified, and publishing the information so that it became a useable resource. These publications, termed the Biological Records, were published by the Cave Research Group of Great Britain and later by the BCRA. The 16 parts contain a 38-year dataset, the accumulated work of over a hundred individuals during thousands of person-hours of work, often in uncomfortable conditions. The final part (1978) saw the retirement of Mary Hazelton as Biological Recorder. A few significant studies were carried out in the post-Hazelton era, but they were the exception, rather than the rule.
Biological studies were re-started in 1998, with the work of Paul Wood and John Gunn in the Limestone Research Group at the University of Huddersfield. In 2000 I took over the post of Biological Recorder within BCRA from Mick Day. When I took over, the largest single job remaining from the Hazelton period was the creation of a computer-readable (digital) version of the Biological Records. With financial assistance from BCRA this job was completed in 2003. The digital version is named Hazelton in honour of its main architect. The paper by Proudlove et al. in this issue is the first to use the Hazelton dataset (with additional records from elsewhere) to examine the distribution of subterranean organisms at the species level. Previous studies, by Hazelton and Glennie, by Jefferson, and by Chapman, have described the biota principally at the phylum, class and order levels. Further analyses using the Hazelton database are now in progress. A copy of the full Hazelton database will be available on the BCRA biology web page within a few months.
The completion of a digital version of the Biological Records, and its subsequent analysis, sees the culmination of the general data gathering period. Randomly collected data, such as form the core of Hazelton, are no longer required for several reasons: 1, A dataset of >6000 records from a 38 year period is quite adequate, and further similar data would add little, if anything, to knowledge. 2. Although Mary Hazelton managed to get all of the specimens sent to her properly identified this is not so easy today. The experts who can identify difficult animal groups have neither time nor resources to devote to "our" samples. Notwithstanding this problem, it cannot be over-emphasised that accurate identifications are crucial to any sort of biological study. Organisms that are not identified (ideally to species), or, worse, are identified wrongly, have no place in modern biology. Great damage would result from any conclusions drawn using misidentified organisms. 3. It is probably true to say that such random collection is contrary to the principles of conservation.
The main conclusion we draw from these data are: Britain and Ireland have very few obligate subterranean animals (troglobites and stygobites) and we hypothesize that this results from the effects of the Pleistocene glaciations. A large proportion of records are of accidentals, those animals that have no place in the subterranean habitat. Another large proportion are troglophiles, which are viable in both surface and subterranean habitats. Some of these will be active colonizers and we can characterize the current subterranean fauna as an early post-glacial one. One intriguing possibility, suggested in the Crustacea data presented later, is that some stygobites may have survived Pleistocene glacials in caves beneath the ice - sub-glacial refugia. We are currently examining this with DNA profiles.
To assist in determining the directions of future work we can use the Hazelton data to pinpoint areas where we are lacking knowledge, and areas that are of obvious importance within the British subterranean realm. We can thus plan and implement targeted studies. For example, it is a surprising fact that terrestrial areas of Peak Cavern are almost unstudied. This large, old and diverse cave would form an ideal site for a whole cave study. Ogof Draenen also comes to mind and is rather better preserved than Peak Cavern. The Hazelton data also demonstrate that Collembola are among the most important of all animals within British caves. These small to minute animals require a determined approach but any study would be very worthwhile, especially as there is currently a major study of this group under way at Reading University. A further insight from Hazelton is that we know quite a lot about caves but rather less about an equally significant subterranean environment, the groundwater. This suite of habitats covers much greater areas than do caves. Very recently one major water company, whose supplies come predominantly from groundwater, have begun active study of the groundwater organisms found in their boreholes. I expect some major advances in knowledge of the groundwater fauna to come from this collaborative project.
As with every aspect of modern life, subterranean biology gets more complex by the year. Although the time of the "Gentleperson-Amateur" is not yet passed, it is quickly receding. Current studies in progress require, among other things, deep borehole sampling, DNA analysis and stable isotope analysis. These are not kitchen-sink activities. I predict that progress in subterranean biology in future will rely very heavily on University or similar research facilities. Of course, this is no different to many other aspects of scientific endeavor.
So to summarize my position on where we go from here: No random collections, no collection of accidentals, collection of targeted taxa only, collections in targeted sites only. It is perhaps too early to ask for hypothesis-driven research and in any case I do not believe that this should entirely supplant observational activities.

Papers

(Proudlove, G.S., P.J. Wood, P.T. Harding, D.J. Horne, T. Gledhill & L.R.F.D. Knight)
A review of the status and distribution of the subterranean aquatic Crustacea of Britain and Ireland.
Vol 30 (2) pp 53 - 74

Abstract: Britain and Ireland do not support many obligate subterranean organisms (troglobites and stygobites) compared to mainland Europe. The largest group of stygobitic taxa is the Amphipoda with seven representatives. In addition, there is one isopod, one syncarid and one copepod. This paper examines the distribution of these animals based on samples taken over 150 years. There is also a number of stygophilic taxa (Copepoda, Cladocera, Ostracoda and Amphipoda) that may be common and even abundant in some caves. Several taxa previously identified as stygobites in Britain and Ireland are shown to be stygophilic. Most stygobitic taxa appear to be restricted to an area south of the maximum limit of the Devensian glaciation. However, there are exceptions, the most significant of which is the presence of a syncarid in central Scotland. Dispersal from southern European refugia, and survival in the glaciated areas in tundra or sub-glacial refugia are discussed. It is concluded that both mechanisms may have played a part in influencing the current distribution of the fauna. Much additional research on these animals is clearly required. In particular, a modern systematic survey of subterranean habitats (hyporheic, hypotelminorheic and deep phreatic groundwaters) and a phylogeographical analysis to examine the relationship of the British and Irish fauna with that of mainland Europe. Dedication: To the memory of Mary Hazelton (1905- 1987) without whose tireless work, as the Biological Recorder of the Cave Research Group of Great Britain, this paper could not have been written.

(Fitzpatrick, S.M.)
An archaeological study on the extraction of flowstone from caves and rockshelters for producing stone money in Western Micronesia.
Vol 30 (2) pp 75 - 82

Abstract: For centuries, Yapese Islanders in western Micronesia traveled 400Km south to the Palauan archipelago to carve their famous stone money from speleothem found in caves and rockshelters. Ethnohistorical accounts and oral traditions provide some information regarding the manufacture of stone money, but a lack of archaeological data has prevented a better understanding of how limestone was quarried, the residuals from carving stone money, and the mineralogical composition of carbonates used in production. In this paper, the first archaeological evidence for speleothem quarrying in Palau by the Yapese is presented. Macroscopic and mineralogical analysis of speleothem debitage and stone money disks indicate that 1) the material quarried was culturally produced and not natural detritus, and 2) it was calcite and not any other material type commonly cited by historians and archaeologists.

(Murphy, P.J. & T. Lord)
Victoria Cave, Yorkshire, UK: new thoughts on an old site.
Vol 30 (2) pp 83 - 88

Abstract: A re-examination of the 'laminated clays' overlying the Ipswichian strata in Victoria Cave suggests an origin as a cave interior deposit rather than a pro-glacial deposit coincidentally located within a cave. These sediments pre-date the second phase of the Late Glacial Imerstadial. The sequence exposed outside the cave mouth shows that the area was glaciated post-dating OIS 5e then subjected to a prolonged period of periglacial weathering pre-dating the Late Glacial Interstadial.

(Sustersic, F., S. Sustersic & U. Stepisnik)
The Late Quaternary dynamics of Planinska jama, south-central Slovenia.
Vol 30 (2) pp 89 - 96

Abstract: During the last l00Ka cave development in Planinska jama has been very "vigorous", but it has not essentially modified the pre-existing layout of bedrock channels. Spatial interpretation of the sediments in Planinska jama has revealed that only the Pivka river can be proven to have flowed through the cave in the Early Quaternary. Its outlet switched between the present cave "entrance" and the Malni Valley. After Würm I the Cerkniscica river entered Cerknisko Polje. Gradual elimination of the polje's main vertical ponors by its growing alluvial cone deflected the main stream westwards. This water found its way to Planinska jama, up to this time penetrated only by the Pivka. Newly arrived waters pushed the Pivka from the Eastern Branch of Planinska Jama and made the river reopen a long-choked outlet in the area of the present cave "entrance" before the end of Würm II. Consequently, the Western and Eastern branches of Planinska jama were washed clear of older sediments and eventually the present flow pattern was established.

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