Foreword top ↑
Why do living organisms have the designs (and especially the
skeletons) that they actually possess? Is it possible, and legitimate,
to infer from the fossilised remains of a long-dead creature how it
functioned as a living system, with all the components operating
together in harmony? Some 40 years ago there was an often stated view
that studies of functional morphology in fossil animals could never be
more than clever speculation. Yet as time went by, it became
increasingly clear that functional interpretations, when carried out
in the right way, were indeed a proper field for study in
palaeontology, and that animal skeletons, of almost any kind, could
yield definitive information about how their bearers had lived.
We need first to consider the origins of animal skeletons. There are
two important factors here. The first is contingency, in other words
the `accidents of history', which established suites of body plans
which could subsequently be modified in different ways. Yet as ROGER
THOMAS and WOLF-ERNST REIF pointed out in their `Skeleton-Space' model
(1993), there are confining physico-chemical constratints which
thereafter determine evolutionary pathways. There are, in fact, only a
limited number of ways in which a skeleton can be functional, as
determined by the properties of the material of which it is
constructed, constraints upon growth and development, and the
requirement for its component parts to function in terms of the whole
organism. In consequence "the discovery of `good' designs those that
are viable and that can be constructed with available materials was
inevitable, and in principle predictable ... the recurring designs we
observed are attractors, orderly and stable configurations of matter
that must necessarily emerge in the course of evolution" (THOMAS &
REIF 1993).
Where then, with this in mind, do we proceed from here? Amongst
compendia regarding form and function in fossils, we have the recent
Functional Morphology of the Invertebrate Skeleton (1999), a fine
collection of 43 papers edited by ENRICO SAVAZZI. Here one finds both
specialised case histories and encompassing reviews, dealing with many
kinds of invertebrate, and very useful it is regarding the various
ways in which invertebrate palaeontologists study their fossils as
living organisms. But the present volume is something different, for
it encapsulates the refreshingly individual approach which has emerged
in Germany over the last several years, most vigorously articulated by
MICHAEL GUDO and his colleagues at the Senckeneberg Institute,
Frankfurt am Main. Their basic concept is that the structural and
functional constraints on living organisms can best be interpreted in
terms of engineering analogues. Mechanical engineering, after all is
about how machines are constructed and how they work, and there are
simple analogues all around us. Consider, for a moment the evident
correspondence between the claw of a crab and a pair of pincers, or an
arthropod limb and the arm of a mechanical digger. There are surely
many useful insights to be derived from an understanding of
engineering principles, and the research papers collected in the
present volume are a testament to the vigour of this approach. For
herein we find not only concepts, but also tools and techniques in
common use in engineering applied to biomechanics; computer-aided
design and tomography, landmark analysis, Finite Element Analysis, and
CAT-scans. Such tools give a much greater objectivity to analysis of
function, for it is true enough, as Carpenter comments in this volume,
that `theoretical models are often tainted with preconcieved ideas'.
There are thirty papers in five sections, each of which consists of
several papers, and at the beginning of each section is an explanatory
introduction and summary.
Section 1, Functional morphology and biomechanics.
Following introductory comments by GUDO et al., there
are six papers all concerned with vertebrates, and especially
dinosaurs. To bring up a simple issue, how can we be sure about the
stance of dinosaurs? We need to tackle this from the engineering point
of view, and only if we do this can we be fully objective, and thus
understand behavioural specialisations. The work presented here
employs landmark analysis (BASZIO & WEBER), parametric modelling,
allowing 3D digital reconstruction (the DinoMorph model) (STEVENS),
and various other morphometric tools are developed and explained (EGI
& WEISHAMPEL). Actualistic models, supplemented by CAT scans
(CARPENTER) allow a precise understanding of how forelimbs are
specialised in different groups of fossil predators, and Finite
Element Analysis (FEA) is used to determine stress and strain in
tyrannosaur bones, and so enables testing of ligamentary functions,
and skull resistance to bending in other carnivores (SNIVELY &
RUSSELL). Joint and movement functions (as the more elderly of us know
too well) are adversely affected by continued stresses, and the
skeleton must be designed to cope with this. How it does so is
addressed by HENDERSON & WEISHAMPEL. The techniques now available have
seldom been used before, and have a great analytical power for
palaeontology.
Section 2. Functional and Ecological Morphology,
explores the interplay of biological, ecological, and taphonomic
factors in reconstructing extinct organisms. Here again the emphasis
is on vertebrates, both living and fossil. Some very interesting
questions arise, and are answered. Was a bipedal posture possible for
a Triassic lizard (RENESTO et al.)? Yes. Can limb reduction in living
fossorial skinks be quantitatively correlated with burrowing
performance (BENESCH & WITHERS)? Yes. What can be said about the three
dimensional kinematics of climbing in various arboreal apes? Can this
knowledge be applied to the understanding origins of human bipedalism
(ISLER)? Probably. Is the optimisation of molar dentition in small
rodents applicable also to larger animals (SCHMIDT-KITTLER)? No. Are
structural specialisations of arvicoline rodent teeth related to diet
(HERRMANN)? Yes. Are the teeth optimised in Miocene Hipparion for shea
ring (KAISER)? Yes, they are, but without investment in unnecessary
structure.
Questions as these are fully explored in this section, and while the
papers are on the whole less mathematical, they still rely directly on
engineering principles. So do the three papers in this section dealing
with invertebrates. Thus a modern coral living on soft substrates,
swept by tidal action, can passively right itself if overturned, but
only if its shape is precisely constrained (HUBMANN et al.). Both
mathematical and physical models are used here to shed light on the
orientative function of the nema in scandent graptolites (RANTELL &
RIGBY). And encrusting bryozoans grow in different ways according to
whether or not they are covering microbial mats; herein the different
kinds of patterns are quantified (KASELOWSKY et al.).
Section 3. Engineering and constructional morphology discusses
how specific structures work in the context of the whole organism, and
it is here that the use of FEA comes into its own. This means that
structures can be modelled as a set of small, clearly defined elements
interconnected at nodal points, and this has important
applications. While some simplifications inevitably result, the models
produced can be tested and evaluated independently. The use of this
tool is here explained (FASTNACHT et al.), and in separate papers is
applied to various problems. For example, the shapes of reptile and
mammalian skulls relate, very precisely, to stress patterns generated
by biting and supporting the head (PREUSCHOFT & WITZEL). The
application of FEA to carnivorous synapsid reptiles (JENKINS) provides
clear indications about comparative predatory habits, and hence niche
partitioning. Likewise, the detailed analysis of the head of a dogfish
(GUDO & HOMBERGER) provides evidence of mechanical constraints for the
evolutionary history of fishes and tetrapods and bears very directly
on the fish-tertrapod transition. In the final paper in this section
GUTMANN combines a philosophical, constructional, and mathematical
approach to problems of crustacean evolution
He comments "Constructional morphology is a necessary prerequisite of
evolution theory" Who could doubt it?
Section 4. Constructional morphology and evolution applies the
biomechanical approach to major issues in metazoan evolution, with
particular reference to the constraints governing evolutionary
pathways. OSCHMANN et al. set the scene with a consideration of the
early evolution of the Earth and the origin of life. Here is stressed
the importance of the ancestral `motiloids', fluid-filled
membrane-bound protocells, from which came procaryotes and later the
eucaryotes through endosymbiosis. While the ideas of GRASSHOF and GUDO
have already been presented in the Senckenberg Poster The Evolution of
Animals' (2001), here they are spelt out in greater detail. Moreover
the concept of an ancestral `gallertoid' is here explained, as central
to the development of compact bodies on the one hand and elongated
bodies on the other. The nearest living creature to the hypothetical
gallertoid is the placozoan Trichoplax, which is here considered (SYED
& SCHIERWATER) and its evolutionary history reconstructed. Lastly an
engaging and extended article by GUDO treats the evolution of the
chordate Bauplan and its modifications. While some of the theoretic
models here are going to be controversial, they are surely worthy of
attention, and they should be much debated. These evolutionary
pathways are after all dictated by structural and functional
constraints, and their elucidation is founded on an understanding of
engineering and biomechanics what will, and what will not work.
Section 5, Theoretical concepts. ethodology and philosophical
analysis concerns the validity and methological justification of our
current evolutionary framework. The function of the philospher, as
ISIAH BERLIN pointed out many years ago is to question, maybe not too
often, and maybe not too much, the structures and frameworks that
govern our thinking and that we take for granted in our society. So,
is a rigidly prescriptive cladistic framework really the best way
forward for phylogenetic reconstruction? REIF provides another way of
thinking, in terms of `reciprocal illumination' of pattern and
process. He notes "The final goal of all systematic and evolutionary
studies are integrated stories of the adaptive, synecological, and
biogeographic histories of taxa. It does not seem prudent to stop
short and restrict oneself to the construction of cladograms." Well
said indeed! What of the geosphere-biosphere system? Is life truly a
planetary phenomenon (LEVIT & SCHOLZ) is the biosphere itself a
living system of the highest order, part of an uninterrupted global
morphospace? With such thoughts we end this remarkable collection of
papers, so remarkably broad-ranging, but so germane to the main
theme. I hope that there will be further papers of similar kind
forthcoming in Senckenbergiana Lethaea. We have here truly a feast
for the intellect. This compendium, for which we thank MICHAEL GUDO
and his colleagues, will surely have a singular influence. I
personally have learned immeasurably from it. I trust that it will
stimulate other readers in the same way.