Collins New Naturalist Library. Philip Chapman

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provided with candles, descended the sloping passage and ladder which led to the depths below.” They spent two hours searching for cave animals and, although they failed to reach the underground river, made a reasonable collection of fauna, including the rare blind cave spider now known as Porrhomma rosenhaueri. In the same year, pioneering English arachnologist F.O.P. Cambridge collected spiders in Wookey Hole, but without finding anything of particular interest.

      Early in 1895 E.A. Martel and his wife paid a well-publicised visit to Ireland. The event prompted the Fauna and Flora Committee of the Royal Irish Academy to support H.L. Jameson with a grant “to further investigate cave fauna in Ireland”. He joined the Martels in the Enniskillen area of Co. Fermanagh and, while the Frenchman surveyed the caves and drew up his plans, Jameson collected cave animals. The interest seems to have persisted, for Jameson is also known to have made faunal collections in Speedwell Mine in Derbyshire in 1901, but there then followed a gap of over thirty years during which British cave fauna was again neglected.

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      Fig. 1.9 One of the earliest illustrations of cave fauna from Adolf Schmidl’s Die Grotten und Höhlen von Adelsberg, Lueg, Planina und Laas. Wien, Braumüller, 1854. (Courtesy Trevor Shaw)

      In 1936 the British Speleological Association was launched, with a brief to co-ordinate the work of caving clubs and to foster interest in the scientific aspects of caving. Things did not run entirely smoothly, however, and in 1947 another body, the Cave Research Group of Great Britain, emerged with a more specific research interest. Both societies ran in parallel until 1973 when they merged to form the British Cave Research Association which has become a major publisher of speleological research.

      Meanwhile another organization concerned with cave science had been formed in 1962. This was the Association of the Pengelly Cave Research Centre, now the William Pengelly Cave Studies Trust. It is London-based, but its interests are very much centred in Devon where it runs the Pengelly Cave Research Centre at Buckfastleigh. The trust is active in education and conservation and produces publications covering a broad range of speleological topics.

      The multidisciplinary nature of speleology allows significant contributions to be made as much by talented amateur observers as by trained professional scientists, and we owe much of our present knowledge of the faunas of British caves to the work of a handful of exceptionally dedicated amateur naturalists. The central figures of the group were Brigadier E.A. (Aubrey) Glennie and his niece Mary Hazelton, who in 1938 began making systematic collections in the caves of Yorkshire, Derbyshire and Mendip. Glennie, an excellent all-round naturalist, picked up his interest in cave life while serving in India, where among other things, he published a study on the nesting behaviour of Himalayan Swiftlets in caves. On his retirement in 1946, he became a driving force in the biological work of the newly formed Cave Research Group, and was soon recognized as an authority on British hypogean amphipods. Hazelton assumed the mantle of Biological Recorder to the CRG, and for the next 29 years diligently co-ordinated the identification of collections submitted by fellow cavers and compiled the results for publication, first in the Transactions of the Cave Research Group and later of the British Cave Research Association. Among the most notable contributors to the faunal collections of this period were Jean Dixon of the Northern Cavern & Mine Research Society and W.G.R. Maxwell of Chelsea Speleological Society.

      The 1950s saw the appearance on the scene of two particularly influential figures, both professional biologists. One was Dr Anne Mason-Williams, a microbiologist whose pioneering studies on the microflora of South Wales caves remains the definitive work in this field. The other was Dr G.T. ‘Jeff’ Jefferson, a lecturer in zoology at University College, Cardiff, who quickly established himself as the leading authority on British cave faunas and went on to become president of the British Cave Research Association, and a greatly respected ambassador for speleology in Britain. Jefferson’s major contribution to cave science in Britain before his untimely death in 1986, was in shaping the wealth of observation gathered by his amateur predecessors into a coherent picture of the biogeographical history and ecological relationships of our cave fauna. It is his work above all that has provided the inspiration for this book.

      Non-cavers are fond of asking cavers why they venture underground. The usual answer is along the lines that “caving is good fun”. Many would add that caving is most fun when spiced with the excitement of discovery. For the sporting caver, this means finding a way into previously unvisited passages, or whole new cave systems. For the speleologist there is the further excitement of recording new observations and of gaining fresh insights into the history, development, or life of the cave. The discipline of cave biology remains poorly developed in Britain and Ireland, affording tremendous scope for discoveries of all sorts by amateur as well as professional naturalists.

      Driving curiosity and a sense of wonder are perhaps the two features which above all unite the caver and the naturalist. I hope that this book can make the passion of the one intelligible to the other, and so enhance the experience of both.

       The Cave Habitat

      Put this question to any wetsuit-clad, hard-hatted individual found walking across the Mendip Hills, Yorkshire Dales or the shining limestone pavements of the Burren, and you will discover that a cave is a naturally-formed hole in limestone which is large enough to be explored by a caver.

      Ask the same caver what he or she has noticed in the way of living creatures in caves, and the answer may well be “not a lot.” It will perhaps surprise most cavers (and naturalists) to learn that over a hundred species of invertebrate animal have been recorded as maintaining permanent populations in cave habitats in Britain and Ireland, plus another score or so species of creatures such as bats and moths which use caves as a regular part-time shelter. The cryptic community to which these creatures belong remains largely undetected by cavers because it generally avoids the relatively large tunnels and booming chambers, the glistening calcite draperies and crashing waterfalls which so captivate the human cave enthusiast, preferring instead the cosy confines of smaller cracks and crevices. I hope in this book to shed light on at least a portion of this unsuspected world which lies beneath our feet, whether we live in Grassington or Glasgow, Lisdoonvarna or London, and to offer pointers to fellow amateur naturalists towards fruitful areas of investigation for the future.

      To unravel the natural history of a cave, or indeed any habitat, we must try to perceive it as far as possible from the point of view of its inhabitants. It requires a considerable effort of imagination for such sight-dependent creatures as ourselves to grasp the essence of life in the dark and labyrinthine realm of the cavernicole. A little inspired speculation may be needed to find the ‘right’ questions to lead us to fresh insights into the mysterious world beneath us.

      We might start by imagining what kinds of habitats could be accessible to the cavernicole and then consider which environmental characteristics of such places are likely to influence its choice of where to live. Almost at once we run into problems, for although a good deal is known about the environment in man-sized, air-filled limestone caves in Britain and Ireland, we know much less about the conditions within smaller cracks and crevices or in caves beneath the water-table, and less still about our submarine caves. Fortunately, such information is more readily available from other parts of the world. So in this chapter we will take an international approach to defining and classifying the cave environment, before turning in later chapters to a detailed consideration of what is known about our own cave fauna.

      What then is a ‘cave’ as perceived by its inhabitants? The dictionary definition of “a natural underground chamber”

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