Paleobiology: Morphology in the contemporary landscape of syste... http://findarticles.com/p/articles/mi_qa4067/is_200304/ai_n920...

FindArticles > Paleobiology > S p r i n g 2 0 0 3 > Article > Print friendly

Morphology in the contemporary landscape of systematics: Learning from the trilobite

O'Leary, Maureen A

Morphology in the contemporary landscape of systematics: learning from the trilobite

Fossils, Phylogeny, and Form: An Analytical Approach. Edited by Jonathan M. Adrain, Gregory D. Edgecombe, and Bruce S. Lieberman. Kluwer
Academic/Plenum Press, New York. 2001. 402 pages. Cloth $130.00.

Morphological Characters and Contemporary Systematics

Circumscribing morphological characters in contemporary systematics research, and doing this with efficiency on a large scale, is a topic of
paramount importance to practicing systematists. The enormous infusion of data from the relatively young field of molecular systematics has
transformed the entire landscape of systematics research because thousands of molecular sequence characters can be generated quickly,
efficiently, and with little ambiguity. This facilitates the rapid growth of enormous molecular sequence data matrices. Although most
morphological systematists employ modern methods of phylogenetic analysis, the mechanics of collecting morphological data (which present a
variety of complex problems) have not been transformed to keep pace with the developments in molecular systematics. Morphological data
remain difficult to collect, hard to convert to discrete characters, and poorly archived. There remains, for example, no morphological equivalent
of GenBank, the public-access molecular sequence database that systematists use to deposit and retrieve raw data. Without equivalent
infrastructural support, many morphological systematists find themselves recollecting data that have been published only as text descriptions
and often publishing their own work without thoroughly documenting their characters through labeled illustrations. Morphological systematics
thereby advances relatively slowly.

That morphological data (and by this I mean any phenotypic character) should be included in phylogenetic analysis is not in serious dispute. The
need to include such data is subsumed by arguments that more data are better than fewer data (Kluge 1989; Nixon and Carpenter 1996) when
reconstructing phylogeny, and each year combined analyses of fossils, extant taxa, and molecular and phenotypic data appear. However, until
morphological matrices can come to include the hundreds or thousands of characters necessary to describe an organism's phenotype, collected in
an explicit, repeatable fashion (Wiens 2001), morphological systematics risks becoming temporarily marginalized by the onslaught of molecular
sequence data currently being collected for systematics research.

Contemporary Morphological Systematics: Morphometrics to the Rescue?

Do refined measurements (i.e., morphometrics) obviate the problem of turning complex continuous morphological information into discrete
characters for phylogenetic analysis? In Fossils, Phylogeny, and Form: An Analytical Approach I anticipated that most chapters would tackle the
complicated intersection of morphometrics and systematics in problems of both alpha taxonomy and relationships of higher taxa. Instead, the
volume is much more diverse. It addresses these topics in part, but it also devotes significant space to biogeography, stratophylogenetics,
morphological disparity, database construction, and rates of evolution. The volume is unified by investigations of the trilobite-examples from its
taxonomy, systematics, and evolution-but the overview of techniques and methodologies is fully relevant to paleobiologists working on any
taxon. Impressively, as a model organism, the trilobite does not disappoint over such a broad range of topics.

The volume begins with a chapter by McLennan and Brooks, which is a short reprise of information from the 1991 Compleat Cladist (Wiley et al.
1991). The refresher is important because the 1991 book clearly articulates, through definition and examples, many often-neglected cladistic
fundamentals such as Hennig's Auxilliary Principle. Relevant to issues addressed throughout the volume is the following definition: "a character
[italics theirs] is any observable part, or attribute, of an organism" (p. 11). With this definition, McLennan and Brooks foreshadow chapters
dealing with such difficult topics as nondiscrete morphological character data. The beauty of their broad definition is threefold, however: it is
data-inclusive, it is stripped of process assumptions that might result in the a priori elimination of data from phylogenetic analysis, and it
actually captures, for better or worse, how morphological systematists conduct their work every day. However, like a vague law that leaves the
messiness of interpretation to the courts, systematists, particularly morphological systematists, still face a heap of ugly day-to-day decisions
about the identity of a morphological character.

Three chapters in this volume (MacLeod, Zelditch et al., and Hughes and Chapman) deal broadly with the integration of morphometrics and
phylogenetics. Zelditch et al. focus on whether morphometric data are appropriate for phylogenetic analysis at all because some have argued
that owing to the continuous nature of morphometric data, "hypotheses of homology cannot apply to them" (p. 146). Zelditch et al. conclude that
some morphometric methods (e.g., traditional measurements, geometric shape coordinates, and partials warps) are appropriate for phylogenetic
problems because they fit the homology concept like any other character. MacLeod counters in his chapter that all morphometric methods share
certain fundamental similarities that make them equally applicable to phylogenetics. The scope of McLennan and Brooks's definition of character
(noted above), for example, presents no impediment to the use of continuous data. As observed by Wiens (2001), and reiterated by MacLeod, in
practice, the morphological systematics literature abounds with continuous characters, many of which are delineated using no protocol. MacLeod
asks, If the use of continuous characters is already widespread in practice, why should a character be eliminated from consideration simply
because it has been measured?

1 of 4 8/12/2008 12:14 AM
Paleobiology: Morphology in the contemporary landscape of syste... http://findarticles.com/p/articles/mi_qa4067/is_200304/ai_n920...

MacLeod notices, however, that arguments among morphometricians about preferred techniques, such as described in this volume, continue to
happen largely on the sidelines of major systematics research projects, and the persistence of these arguments has done little to advance the role
of morphometrics in phylogenetics. Few of the larger morphological phylogenetic matrices (e.g., >100 characters) rely on morphometrics at all
to discover or delineate the continuous characters they include.

Neither Zelditch et al. nor MacLeod advocates the direct translation of morphometric data into characters and states. Zelditch et al. state that "it
is not possible to generate characters automatically by a purely morphometric analysis" (p. 171), arguing instead that morphometrics has a role
in descriptive anatomy used to aid in "the discovery of characters even if the variables cannot be automatically equated to characters" (p. 193).
MacLeod envisions morphometrics as allowing us to see "discontinuities in systematic datasets that have phylogenetic significance" (p. 226).
However, I am not so sure that this is the case-that our travails of quantitative morphological character data will be obliterated by simply
measuring morphology better, that discrete characters will somehow emerge from what previously seemed nondiscrete. Zelditch et al. recognize
that arbitrary, if explicit, character state boundaries must still be determined by an investigator wishing to turn morphometric variables into
characters.

Finally, Zelditch et al. map their morphometrics-derived characters onto existing trees to investigate how a "homologous feature . . . can be tested
by congruence" (p. 171). If a character can be mapped it can be included in the analysis, which would be a preferable means of testing its impact
on the tree overall.

Hughes and Chapman test the following classic hypothesis: that "inviolable developmental canalization in segment formation" (p. 49) obtains in
certain Silurian trilobites following the initial Cambrian radiations. To do this they map the character-variability of trilobite thoracic
segments-onto a cladogram and show that one trilobite lineage (Aulacopleura konincki) exhibits a pattern of holaspid segmental flexibility yet is
nested within a clade containing basal members with stable holaspid segment numbers. They argue that this result falsifies the notion of
"inviolable" canalization. The test is compelling, although it is unfortunate that the cladogram construction does not appear to be particularly
rigorous (it is not directly generated from the global application of an optimality criterion to all character data). It would also have been
appropriate to include the character, variability in segment number, in the analysis and show its explicit optimization.

Their paper also details extensive morphometric analysis of these Silurian trilobite lineages, to examine differences in degrees of shape variation
among species. They suggest that together these analyses show an "example of how phenetic and phylogenetic approaches can compliment [sic]
one another in providing fresh insights" (p. 52). The morphometric analysis, however, does not appear to be required for the determination of the
number of segments present in the holaspid stage of each species. This can be done by eye, and this is the character that is ultimately examined to
falsify the notion of inviolable developmental canalization. The central insight of their paper comes arguably from phylogenetics alone. Thus
although the authors have framed a test of an important generalization about the Cambrian explosion, it is not clear that the authors have also
forged an alliance between morphometrics and cladistics.

Disparity and Phylogeny

Wills tackles the more phenetic concept of morphological disparity, "differences [italics his] generated by evolution" (p. 57), often studied in some
form of multidimensional space. Many of Wills's comparisons between disparity and phylogeny rely on a characterization of phylogeny as a series
of separate investigations of partitioned character data sets. For example, he argues that as different morphospaces derived from different data
often produce different accounts of disparity, so different cladograms emerge from analyses of different sets of character data. But the latter type
of phylogenetic analysis (data partitioning) has been heavily criticized as a practice that is inconsistent with the logic of parsimony analysis
(Kluge 1989; Nixon and Carpenter 1996), and which should be abandoned in favor of a character congruence approach in which all data are
combined and analyzed simultaneously. That said, perhaps ultimately results of disparity research should remain accountable to comparison
with the "absolute or 'holomorphospace'" (p. 58) that Wills mentions.

One study of morphological disparity in trilobites that Wills cites shows that taxonomic diversity is out of sync with morphological disparity.
Although this seems like a straightforward test and result, it is odd that given that diversity in extinct taxa must be based on morphological
difference, why is it that some morphological diversity is considered important for taxonomic differences and other morphological diversity is
not, short of knowledge of the true species boundaries or the true phylogeny?

Wills notes that the discovery of morphological constraints is a "common goal" (p. 112) of disparity studies, writing that "if an ancient clade fails
to colonize some regions of morphospace after the initial phases of its radiation, it is tempting to speculate that further exploration is constrained"
(p. 112). Until this is framed as a test, however, alternative explanations for the morphospace occupation will remain at large. If this remains a
goal of disparity studies, it is important to articulate how disparity studies can specifically reject the hypothesis of constraint, as has been done
using phylogenetics (e.g., Hughes and Chapman).

Trilobites in Space and Time

Ebach and Edgecombe explore aspects of cladistic biogeography, specifically using methods derived from Component Analysis. By using trilobite
examples they push cladistic biogeography beyond the typical scope because the focus is a marine taxon whose evolutionary history predates the
fragmentation of Pangea. Their examples address the general significance of fossils in biogeographic studies.

Adrain and Westrop present forceful arguments against the relatively recently emerged practices of stratocladistics and "stratolikelihood" (p.
294). They anchor their criticisms in the position that "strato-methods are damned on theoretical grounds alone" (p. 291). This is because

2 of 4 8/12/2008 12:14 AM
Paleobiology: Morphology in the contemporary landscape of syste... http://findarticles.com/p/articles/mi_qa4067/is_200304/ai_n920...

temporal data are not "intrinsic" (p. 295) properties of organisms, making them inappropriate for phylogenetic analysis. They make convincing
arguments that lamentation that morphology alone might be misleading amounts to insufficient justification for incorporating temporal
information into phylogeny reconstruction.

A significant portion of the chapter is devoted to non-theoretical, in other words, practical, problems that may arise with stratomethods. For
example, they argue that the high quality of the stratigraphic record is an assumption that underlies stratophylogenetics and, using a trilobite
example, show the complexities that obtain when this expectation of quality goes unfulfilled. The completeness of the stratigraphic record,
however, may ultimately be no more germane to the argument about whether stratomethods should be used or not than is the anatomical
completeness of a fossil to the usefulness of standard phylogenetic methods. Another tricky angle on this debate is their argument that one of the
central problems with stratigraphie data (and by extension its use in stratocladistics) is that time is not a discrete variable. As other chapters in
the volume discuss, this is not a characteristic unique to temporal data but is also characteristic of many morphological characters.

Adrain and Westrop have, however, primarily focused the discussion where it needs to be: on the issue of intrinsic versus extrinsic characters.
How such characters are defined and what relationships they have to evaluating evolutionary hypotheses remain the critical philosophical
issues at stake in this debate.

Two chapters (Lieberman, Eldredge) are devoted to rates of evolution and the origin of new taxa. Lieberman discusses the study of speciation rates
(in both probabilistic and deterministic frameworks), which he argues has been greatly enhanced by the increase in availability of species-level
phylogenies for fossil taxa. Eldredge writes of the significance of the study of "cross-genealogical patterns" (p. 345) in the history of life, suggesting
that such study might contribute to "an accurate causal theory of evolution" (p. 341). He reflects that the early formulation of punctuated
equilibrium focused on monophyletic taxa but "no attention whatsoever was paid to the (in retrospect, blindingly obvious) fact that many
sympatric and synchronie species of wholly unrelated taxa demonstrate exactly equivalent, and nearly synchronous, patterns of . . . change." (p.
345). He predicts that regional and global environmental disruptions are tightly linked to evolution and that case studies will reveal many
synchronic patterns of clade diversification. Much of his chapter includes further attempts to corroborate punctuated equilibrium (with trilobite
examples).

An obstacle to such cross-genealogical work, Eldredge states, is that scientists have exhibited "legitimate reluctance" (p. 369) to stray outside
their area of taxonomic expertise to comment on patterns of evolution in taxa that co-occur with the ones they have directly studied. This might
explain why Eldredge, when discussing biostratigraphy and the Bighorn Basin, only cites papers that are over 25 years out of date (Gingerich
1976). Ongoing projects in the Early Tertiary of the Bighorn Basin (e.g., Rose and Bown 1984; Wing et al. 1991; Bown et al. 1994; Bloch and
Gingerich 1998) continue to generate important primary data that contribute to the investigation of the kinds of cross-genealogical speciation
patterns and environmental change that Eldredge advocates. Furthermore, that a Bighorn Basin study alone fails to encompass the entire
geographic range of a taxon of interest does not necessarily mean that the pattern that study finds will be refuted should biostratigraphic data
become available for a species-wide study. Finally, the overall argument that an accumulation of such pattern studies will allow us to address
causality or "what drives evolutionary process?" (p. 342) remains underdeveloped in the chapter as a whole.

Finally, Kaesler et al. discuss how the future of paleontology will be greatly aided by the development of useful relational databases. They
describe PaleoBank, a database that draws heavily on the Treatise on Invertebrate Paleontology and manages various types of taxonomic,
anatomical, and geological data. They emphasize that there are often important differences between databases designed to serve collections
management at major museums and those devoted to research tasks, in other words, designed to "manage knowledge" (p. 389). They advocate
that, whenever possible, databases should be modeled in such a way that they can talk to other databases should the need arise in the future.
Because the number of highly trained systematists is dwindling, Kaesler et al. emphasize that there is some urgency to the need to develop
effective archives lest we risk losing extensive knowledge with the passing of various experts.

Keeping Track of Morphology

Although diverse in scope, this volume remains focused on a variety of complicated topics related to morphological systematics. Some of the
chapters could have been more severely edited (several exceed 50 pages), but the overall vision of using trilobites as a general study case for
morphological evolution is compelling.

The systematics of trilobites, a wholly extinct clade nested within Arthropode, the most speciose clade of extant taxa on the planet, lies right at
the intersection of problems of integrating molecular sequence data and morphology by using character congruence (Giribet et al. 2002).
Placing trilobites in future combined analyses makes them an excellent test case for issues of the importance of fossils in phylogeny
reconstruction, the role of missing data, and the use of phylogenetics, in particular, optimization to reconstruct the developmental history of
extinct groups.

The character congruence approach argues for data combination, or the construction of supermatrices, but how are we preparing to keep track of
the data that will be amassed for supermatrix analyses? Will morphologists recollect data over and over before some centralized clearinghouse for
morphological character data emerges?

As mentioned at the outset, the molecular systematics revolution emerged in concert with extensive database support in the form of GenBank.
Even though molecular sequences on GenBank are unaligned and therefore not immediately ready for phylogenetic analysis, all raw data can be
downloaded from the web, a feature that greatly facilitates the ongoing construction of ever-larger molecular sequence data matrices. Similar

3 of 4 8/12/2008 12:14 AM
Paleobiology: Morphology in the contemporary landscape of syste... http://findarticles.com/p/articles/mi_qa4067/is_200304/ai_n920...

infrastructural support, such as a database designed to keep track of images that document morphological observations in systematics, will be
complicated to develop but is becoming increasingly essential for systematics research in general. Just as molecular biologists would no more
discard forever the sequence of letters that describe a gene, morphologists should record what they observe in labeled images so as to document
their own work and to save needless duplication of research.

Literature Cited

Bloch, J. I., and P. D. Gingerich. 1998. Carpolestes simpsoni, new species (Mammalia, Proprimates) from the Late Paleocene of the Clarks Fork
Basin, Wyoming. Contributions from the Museum of Paleontology, University of Michigan 30:131-162.

Bown, T. M., P. A. Holyroyd, and K. D. Rose. 1994. Mammal extinctions, body size, and paleotemperature. Proceedings of the National Academy
of Sciences USA 91:10403-10406.

Gingerich, P. D. 1976. Paleontology and phylogeny: patterns of evolution at the species level in Early Tertiary mammals. American Journal of
Science 276:1-28.

Giribet, G., G. D. Edgecombe, W. C. Wheeler, and C. Babbit. 2002. Phylogeny and systematic position of Opiliones: a combined analysis of
chelicerate relationships using morphological and molecular data. Cladistics 18:5-70.

Kluge, A. G. 1989. A concern for evidence and a phylogenetic hypothesis of relationships among Epicrates (Boidae, Serpentes). Systematic
Zoology 38:7-25.

Nixon, K. C., and J. M. Carpenter. 1996. On simultaneous analysis. Cladistics 12:221-241.

Rose, K. D., and T. M. Bown. 1984. Gradual phyletic evolution at the generic level in early Eocene primates. Nature 309:250-252.

Wiens, J. J. 2001. Character analysis in morphological phylogenetics: problems and solutions. Systematic Biology 50:689-699.

Wiley, E. O., D. Siegel-Causey, D. R. Brooks, and V. A. Funk. 1991. The compleat cladist: a primer of phylogenetic procedures. Special
Publications, University of Kansas Natural History Museum, Lawrence.

Wing, S. L., T. M. Bown, and J. D. Obradovich. 1991. Early Eocene biotic and climatic change in interior western North America. Geology
19:1189-1192.

Maureen A. O'Leary

Department of Anatomical Sciences

HSC T-S (040)

Stony Brook University

Stony Brook, New York 11766

E-mail: moleary@mail.som.sunysb.edu

Copyright Paleontological Society Spring 2003

Provided by ProQuest Information and Learning Company. All rights Reserved

4 of 4 8/12/2008 12:14 AM