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Trilobite paleobiology: Past, present, and future

Adrain, Jonathan M

THE FIRST major international trilobite conference (which also dealt with related arthropod groups) was convened by David Bruton in Oslo in
1973. That meeting resulted in a 467 page proceedings volume (Martinsson, 1975) that has become an indispensible reference on arthropod
paleobiology. With its abundance of high-quality empirical work and influential ideas, the ongoing utility of "Fossils and Strata 4" is assured, yet
it is also a fascinating microcosm of the themes and concerns of paleontology as a whole a quarter of a century ago. Paleoecology blossomed in the
late 1960s and early 1970s. Autecology, and particularly functional morphology, dominates the volume, accounting for about one-third of the
contributions. Plate tectonics was a relatively recent development in 1973 and, following the lead of Whittington and Hughes (1972), several
papers tackled trilobite paleobiogeography. They vary from a bold attempt to determine oceanic circulation through the Ordovician by Ross to
more conventional studies of biogeographic distribution that were aimed at providing further support for concepts of continental drift. There was
a burgeoning interest in community paleoecology, and Richard Fortey's (1975) now-classic paper on the Early Ordovician faunas of Spitsbergen
was the first to define trilobite "communities." The volume was rounded out by diverse papers on trilobite anatomy based on material from the
Burgess Shale and Beecher's Trilobite Bed, the microstructure of the trilobite cuticle, trilobites eyes, high level classification of trilobites, and
various evolutionary themes (character displacement in agnostoids; adaptive radiation in Cambrian trilobites).

The Oslo meeting and its proceedings were by every indication a tremendous success. It is somewhat puzzling, then, that it spawned no sequel. A
glance at any volume of the Journal of Paleontology published in the 1990s will confirm that trilobite study still ranks among the most active
fields of invertebrate paleontology. Yet while workers on other groups (e.g., brachiopods, conodonts, graptolites) schedule regular conferences to
bring their communities together for an exchange of new results and ideas, there have been no specialist meetings devoted to trilobites and
related arthropods since Oslo. The time for such an event seemed ripe, and in August of 1997 we co-organized the Second International Trilobite
Conference, which took place on the campus of Brock University, St. Catharines, Ontario, Canada. Eighty-five trilobite specialists from around
the world gathered for a four day program of talks. In this issue it has been our welcome task to assemble 14 full-length papers derived from those
presentations. The content of this issue and a survey of recent papers published elsewhere provide an opportunity to compare current research
themes with those of 25 years ago, to consider how they reflect the emergence of new areas of interest in the intervening period, and to look
toward the next such meeting, to be held in Oxford in 2001.

Easily the most profound change in trilobite paleobiology, and indeed in systematic biology in general, during the past 25 years has been the
emergence and general acceptance of cladistic methodology. At the time of the Oslo meeting, Niles Eldredge (1972, 1973) was the sole trilobite
worker to embrace the nascent field of phylogenetic systematics. Nowadays, while still far from routine, cladistic parsimony analyses are
commonplace (Adrain, 1998; Adrain and Chatterton, 1994; Edgecombe and Ramskold, 1996; Hughes and Rushton, 1990; Lesperance and
Desbiens, 1995; Lieberman, 1998; Ramskold and Chatterton, 1991; Sundberg and McCollum, 1997). Cladistic approaches have been successfully
applied to longstanding problems ranging from the species-level phylogenies of notoriously intractible families (Adrain and Edgecombe, 1997;
Ramskold and Werdelin, 1991; Westrop et al., 1996) to the high-level classification of major trilobite groups (Fortey and Chatterton, 1988).
Herein, Edgecombe and Ramskold use the spectacular Early Cambrian fossils from China's Chengjiang lagerstatte in a cladistic analysis of
perhaps the most basic phylogenetic issue in trilobite study-the systematic position of the trilobite clade and basal arachnomorph relationships.

As interest in the hierarchical relationships of taxa and the distribution of characters grew, so too did one of the richest sources of phylogenetic
data: silicified larval and post-larval ontogenies. Because trilobite hardparts are calcified from an early stage in larval development, it is often
possible to assemble nearly complete growth series for the entire exoskeleton. Harry Whittington's classic work (e.g., 1956, 1959) revealed the
potential of silicified faunas. Fortey and Owens's (1975) proposal of the order Proetida was among the first studies to incorporate comparative
ontogenetic data into phylogenetic analyses, an approach that has continued to lead to fresh insights into the relationships of taxa (Chatterton et
al., 1990, 1994). In the past quarter century, Brian Chatterton has taken study of trilobite ontogenies to a new level of understanding (e.g.,
Chatterton, 1971, 1980; Chatterton and Speyer, 1997). With increases in SEM technology and a growing appreciation that silicification is a
fairly common phenomenon, ontogenetic work has the potential to unlock many of the remaining long-standing problems of trilobite
classification and evolution. In this issue, Chatterton and colleagues illustrate the first fully documented ontogenies of the family Telephinidae
and in the process raise important issues about the composition and ancestry of the order Proetida. Cr*nier et al. contribute a beautifully
described growth series of a Late Devonian phacopid, a group for which early ontogenetic data has been scarce, despite its cosmopolitan
distribution.

Paleobiogeography, a major concern of the previous generation of trilobite workers, has apparently lost its luster. This decline in interest perhaps
reflects the fact that plate tectonics and continental drift are now core organizing principles of the earth sciences rather than novel hypotheses to
be evaluated with faunal data. Cladistic biogeography certainly offers new approaches to problems such as speciation but with few exceptions
(e.g., Lieberman, 1997), this avenue of research has yet to be exploited.

Paleoecology is as important as ever but the thrust of research has changed. Functional morphology, one of the core areas of trilobite paleoecology
25 years ago, has ceased to be pursued in any meaningful way. Studies of trilobite "communities" have flourished, but the emphasis has shifted
from simple description to consideration of their macroevolutionary or biostratigraphic significance. This change in focus can be traced to the rise

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of evolutionary paleobiology, a movement that has led to a wholesale transformation of how paleontologists view their discipline (the inaugural
issue of the journal Paleobiology appeared a few months before the publication of the Martinsson volume). As early as 1980 Gould had noted that
classical functional morphology and descriptive community paleoecology were waning because they failed to address questions of evolutionary
significance. Five years later, "evolutionary paleoecology" had become firmly entrenched in the lexicon of paleontology (Kitchell, 1985; see also
Valentine, 1973) and is now a flourishing discipline in its own right. Paleoecological studies of trilobites have, and will continue, to contribute to
such areas as mass extinctions (Ludvigsen and Westrop, 1983; Westrop, 1989; Chatterton and Speyer, 1989; Owen et al., 1991), evolutionary
radiations (Fortey and Owens, 1990; Adrain et al., 1998), turnover rates and coordinated stasis in paleocommunities (Fortey, 1980; Westrop,
1996), and patterns and processes of major biotic transition in the history of life (Westrop et al., 1995; Westrop and Adrain, 1998). Herein,
evolutionary paleobiology forms the single largest category of papers in the issue, including studies of trilobite diversity and distribution
(Brezinski), turnover and extinctions (Mikulic and Kluessendorf; Westrop and Cuggy), biostratigraphy and biofacies (Taylor et al.), and
Cambrian macroevolution (Lieberman).

The transformation of paleoecology has been parallelled by a resurgent interest in the ways in which fossil accumulations are formed. At the time
of the last international trilobite conference, many participants would have been hard-pressed to define taphonomy, which, if discussed at all,
was viewed as an area concerned with the limitations of paleontology (e.g., Lawrence, 1968). Taphonomy has emerged as a vibrant discipline
that has stepped beyond this narrow vision (Behrensmeyer and Kidwell, 1985). Trilobite workers have been at the forefront of this new
taphonomic research and have demonstrated novel applications of preservational data in paleoecology (Speyer and Brett, 1985) and in
sedimentary facies analysis (Speyer and Brett, 1986). More conventional work on the interpretation of trilobite accumulations has also
flourished, with contributions ranging from experimental (Hesselbo, 1987; Lask, 1993) and field-based (Speyer, 1987) studies of sclerite
orientation and its hydrodynamic significance, to the implications of taphonomic sorting for paleoecologic interpretation (Westrop, 1986;
Mikulic, 1990). This issue features detailed taphonomic studies of two Late Ordovician trilobite assemblages: Hughes and Cooper look at a
calymenid-dominated deposit from the Kope Formation of Ohio, while Brett and colleagues examine the famous Walcott-Rust Quarry of New York
State along with providing a much-needed modern overview of the fauna.

Morphometric analysis of trilobites was completely absent from the Martinsson volume (with the exception of Temple's plea for a standardization
of trilobite measurements), but has become a growth industry in recent years, with contributions ranging from comprehensive surveys of the
group as a whole (Foote, 1991, 1993), to detailed investigations of a single species (Hughes, 1994; Hughes and Chapman, 1995). The expansion
in morphometric approaches must stem in part from the advent of fast computers and software packages that make even the most arcane
multivariate technique readily accessible. However, it also reflects the increased interest in evolutionary problems such as patterns of
morphospace occupancy over time and the evolutionary significance of morphologic variability. Detailed morphometric analyses are carried out
by Webster and Hughes on the marked distortions introduced by compaction preservation, even when the preservation of detail is very fine, and
by McCormick and Fortey in an attempt to prove the species identity of widespread geographic occurences of a pelagic Ordovician trilobite.

Two papers reflect a continuity of scholarship rather than the departures of the past two decades. Dalingwater et al. continue their work on the
exquisitely preserved ultrastructure of Silurian trilobites, a series that extends naturally from the senior author's papers in the Martinsson
volume. Fortey and Droser, using fieldbased descriptive paleontology, show that trilobite biostratigraphy and systematics have much to
contribute to debates about major international stratigraphic boundaries, with their unexpected finds immediately above the
Ibexian-Whiterockian boundary in the Great Basin.

Given the changes in emphasis in trilobite studies over the past quarter-century, looking ahead even a few years is a risky business. In 1975
nobody would have predicted the decline of functional morphology and paleobiogeography. Based on current trends, cladistics is likely to remain
the method of choice in systematics and phylogeny reconstruction. We expect increasing numbers of trilobite workers to incorporate
morphometric techniques into their studies, and we would be surprised indeed if the current emphasis on evolutionary paleoecology and
taphonomy became reversed in the foreseeable future. However, we would also be surprised if the future did not bring some new developments,
and the Oxford Conference of 2001 will provide another opportunity to guage current trends in trilobite paleobiology. ACKNOWLEDGMENTS

We are very grateful to the Paleontological Society Council for the opportunity to publish these papers as an issue of the Journal, and to D. B.
Blake, former Managing Editor, for his help with the proposal.

Compiling an entire issue of trilobite papers has meant enlisting the aid of a significant proportion of the international community. We are
grateful for constructive and timely reviews from all involved, including P A. Allison, R. Bambach, U. Brand, C. E. Brett, D. R. Brooks, D. L.
Bruton, R. Chapman, B. D. E. Chatterton, E. N. K. Clarkson, G. D. Edgecombe, R. Feist, S. L. Finney, M. Foote, R. A. Fortey, N. C. Hughes, M. E.
Johnson, E. Landing, B. S. Lieberman, J. D. Loch, R. Ludvigsen, K. J. MacNamara, R. M. Owens, A. R. Palmer, B. R. Pratt, R. A. Robison, A. W. A.
Rushton, F C. Shaw, H. B. Whittington, M. A. Wills, and Anonymous. The Second International Trilobite Conference was made possible through
organizational support from D. M. Rudkin, M. B. Cuggy, B. Mills, W. Laslo and the staff at Conference Services, Brock University. Field trips were
organized by D. Armstrong, K. D. Brett, Chatterton, Landing, Pratt, Rudkin and Westrop. Financial support was received from the Brock
University Conference Speaker Fund and the Institute for Cambrian Studies. B. Levman secured corporate funding from a variety of sources.

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JONATHAN M. ADRAIN AND STEPHEN R. WESTROP

Department of Palaeontology, The Natural History Museum, Cromwell Road, London SW7 5BD, United Kingdom,

Oklahoma Museum of Natural History and School of Geology and Geophysics, University of Oklahoma, Norman, OK 73019,

Copyright Paleontological Society Mar 1999

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