Squeezing the Fossil Record

FEATURE fossils up into the chalk country, he had recognized at Although uncertain about the mechanisms in- least part of the stratigraphic column that was later volved, Whitehurst argued for the phenomenon of mapped by William Smith. igneous intrusion as well as extrusion. He recognized that mineralization led to veins cutting limestones; and he knew something of the nature of faulting, Whitehurst’s attainments even if confused by the alleged gulfs under the River In the Inquiry, Whitehurst used observation and ap- Derwent. With the lack of any Whitehurst archives or plied the laws of nature to deduce several basic prin- of records of the Lunar Society it is difficult to say how ciples of geology: much of his ideas came from discussions with others, 1 The Earth had a history of development from a and similarly difficult to say to what extent he influ- chaotic fluid state to one of regular order. enced the ideas of his successors, although White 2 This history was deduced from observations Watson and Farey both followed his Matlock which led to recognition of the principle of stratigraphy. From the few records that survive, it superposition. seems that an outline of the Inquiry was already in his 3 A regular sequence of strata had economic mind by 1763, 15 years before publication, and he implications – i.e. prediction of what might occur had expanded his ideas as a result of Ferber’s visit in elsewhere. 1769. Even then it took another nine years before it 4 Coal originated from ancient vegetation, whilst was completed. If Whitehurst had not been distracted limestone contained marine shells. by his clock-making business and later his duties as 5 Volcanic activity had occurred just as much, or Stamper of the Money Weights, he might have left us even more, in the past as in recent historical times. a legacy of the basis of geological science a few years 6 Certain strata were distinguishable by their fossil before the recognized ‘giants’ James Hutton and contents. William Smith. 7 The sequence of Carboniferous strata around Matlock could be recognized elsewhere. Suggestions for further reading 8 ‘Convulsions’ explained the contorted strata of mountainous regions. Challinor, J. 1947. From Whitehurst’s Inquiry to 9 The disposition of strata could be illustrated by Farey’s Derbyshire: a chapter in the history of Eng- means of sections. lish geology, Transactions of the North Staffordshire 10 The Coal Measure fossil Carbonicola could be Field Club, v.81, pp.52–88. compared with the modern freshwater mussel Unio. Craven, M. 1996. John Whitehurst of Derby, clockmaker 11 The abundant fossil stems in the Derbyshire and scientist, 1713–1788, published by Mayfield limestones were equivalent to the crinoids recently Books, Mayfield, Ashbourne, Derbyshire. 272pp. found in the Caribbean. 12 Heat affected the rocks beneath an intrusion – i.e. contact metamorphism. Topics Cornelis J Veltkamp* and Squeezing the fossil record: unravelling the Stephen K Donovan† evolution of stalked crinoids in the *School of Biological Sciences, University of Liverpool, PO Box 147, Caribbean Liverpool L69 3BX, UK †Department of Palaeontology, Nationaal The fossil record has often been tainted by the label of What is relevant is not whether the fossil record is Natuurhistorisch Museum, ‘incomplete’, but such an insignia might be attached complete but, rather, is it adequate to answer ques- Postbus 9517, NL-2300 RA to our knowledge of any of the natural sciences. tions regarding the history of life? Available evidence Leiden, The Netherlands © Blackwell Science Ltd, Geology Today, Vol. 18, No. 3, May–June 2002 107 Untitled-1 107 09/08/02, 14:40 TOPICS overwhelmingly indicates that it is. However, an ‘ad- for this is that for deep-water sedimentary rocks to be equate’ fossil record may nevertheless present prob- exposed at the modern surface of the land, they have lems of interpretation. Many organisms have a to be uplifted by considerable tectonic forces that only multicomponent skeleton that disarticulates into its act in certain areas of the globe. One such place is the component parts, partly during life in some groups Caribbean. Approximately coeval sedimentary depos- (e.g. arthropods, vascular plants), but completely in its exposed on three widely separated Caribbean is- all such organisms following death (e.g. vertebrates). lands, each formed in rather different palaeoenviron- Thus an oak tree may generate very numerous mental and tectonic settings, have now yielded leaves, flowers, pollen and acorns during life, but faunas of disarticulated sea-lily plates (called ossicles). most of its woody tissues – roots, trunk, branches and This is the first time that fossil crinoids have been twigs – only become available for fossilization after found from a restricted time interval on more than death. An analogous situation among animals is the one Antillean island, and enables authoritative com- numerous moult stages in the life cycles of arthro- ment to be made on their distribution and evolution pods. for the first time. As Cenozoic fossil crinoids have a Other groups contribute most of their skeletal ele- very patchy fossil record globally, these results are of ments to the fossil record after death, such as chitons, more than just local interest. the vertebrates and – the subject of this article – the echinoderms. Most of the fragmentary remains of or- The Miocene Antillean crinoids ganisms with multicomponent skeletons are small or minute, and the SEM is now used routinely to ob- Deep-water deposits containing sea lilies in the Lower serve, interpret and illustrate their features, from fos- and Middle Miocene (about 23.5–11 million years sil pollen and spores through the juvenile moult old) of the Antilles are known from Jamaica in the stages of extinct arthropods to rodent teeth. Our inter- west and Barbados and the tiny Grenadine island of pretations of Caribbean fossil echinoderms, including Carriacou in the east. The Jamaican deposits (Lower the results discussed herein, have relied heavily on Miocene), exposed in a disused quarry near the north the Philips XL30 scanning electron microscope at the coast, are chalks that were laid down in at least University of Liverpool, which has enabled examina- 500 m of water, based on evidence provided by the tion of both the gross morphological features (Fig. 1) enclosed fossil fishes. Crinoid remains have proved to and, where apparent, the stereom microstructure be particularly rare and less than 10 ossicles have (e.g. Fig. 1e), providing exquisite images for system- been found in 15 years of collecting. atic descriptions and comparison between localities. In Barbados, the Lower Miocene crinoid-bearing In modern seas there are five groups of beds form part of the accretionary prism – i.e. the echinoderm – the echinoids (sea urchins), holothur- wedge of sediment (mainly muds, silts and sands) sev- oids (sea cucumbers), asteroids (starfishes), ophi- eral kilometres thick that has been scraped off of the uroids (brittlestars) and crinoids (unstalked feather South Atlantic tectonic plate as it has been subducted stars and stalked sea lilies). The last of these, the beneath the Caribbean tectonic plate. These sedimen- stalked sea lilies, are an ancient group of sessile, at- tary rocks were probably deposited at a water depth tached, filter-feeding invertebrates that formed a measurable in kilometres. Crinoid ossicles have been prominent component of the marine benthos during found as rare specimens in the collections of the the Palaeozoic (about 540–250 million years ago). Naturhistorisches Museum Basel, picked from sam- However, the sea lilies almost disappeared in the mass ples taken from sections that are no longer exposed. extinction at the end of the Permian (250 million In Carriacou, the crinoid deposits are sedimentary years ago) and subsequently failed to return to their rocks at the base of a sequence derived from former abundance and diversity. Further, during the volcaniclastic deposits that have been washed into past 100 million years or so they have disappeared deep water from around the ancient volcanoes that from shallow-water environments as certain major formed the island. Deposition occurred in at least groups of predators have increased in numbers and 150–200 m, based on the evidence provided by a va- diversity. riety of deep-water fossils, including isocrinid crinoids This historic pattern has two major implications (see below). Unlike Jamaica and Barbados, the for the post-Palaeozoic fossil record of sea lilies. Their crinoid-bearing deposits on Carriacou contain a huge diversity was greatly reduced from its peak in the abundance of ossicles and are readily accessible, al- Palaeozoic, so they are relatively uncommon fossils in though diversity is limited to only four species. the post-Palaeozoic fossil record. Further, by vacating Thus Antillean Miocene crinoids are known in shallow-water environments, they are limited to sedi- deposits representing environments as disparate as mentary rocks deposited in deeper water, which are deep-water chalks, bathyal or abyssal muds and the not as commonly exposed subaerially as those laid submarine slopes of an extinct volcano. Although down in shallow-marine settings. The simple reason these environments were obviously very different, the 108 © Blackwell Science Ltd, Geology Today, Vol. 18, No. 3, May–June 2002 Untitled-1 108 09/08/02, 14:40 TOPICS crinoids that lived in them were remarkably similar. Although based on somewhat minimal material, it However, no complete crinoids are known from any appears that, like the closely related echinoids, the Antillean island. Most of the Miocene specimens are modern crinoid fauna of the Antilles probably arose preserved as isolated ossicles of the stalk, called after a pattern of extinctions during the Eocene– columnals (Fig. 1a–d), and rarely as short lengths of Oligocene transition. stalk. Ossicles of the arms and roots (Fig. 1e) are also It is also notable that this is more than just a local present, although more difficult to identify to species.

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