Memoir cover 0.tif - Ohio University College of Osteopathic Medicine
Memoir cover 0.tif - Ohio University College of Osteopathic Medicine
Memoir cover 0.tif - Ohio University College of Osteopathic Medicine
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SOCIETY OF VERTEBRATE PALEONTOLOGY, MEMOIR 3<br />
rotation around<br />
outgroup node<br />
Extant Phylogenetic Bracket<br />
FIGURE 2. Inverted Pyramid <strong>of</strong> Inference. Inferences about the s<strong>of</strong>ttissue<br />
relations <strong>of</strong> bony structures are the foundation and justification<br />
for many paleobiological inferences. The pyramid is inverted to highlight<br />
the point that mistakes in s<strong>of</strong>t-tissue inference cascade up the hierarchy,<br />
magnifying the error. (Modified from Witmer, 1992a, 1995a.)<br />
spurious hypotheses about the structure and evolution <strong>of</strong> whole<br />
communities. As we will see, complicated paleoecological and<br />
evolutionary scenarios have been tied explicitly to particular<br />
notions about the s<strong>of</strong>t-tissue relations <strong>of</strong> the antorbital cavity <strong>of</strong><br />
archosaurs. Thus, a seemingly arcane issue can have important-and<br />
unexpected-ramifications (see Witmer, 1992a,<br />
1995a for other examples).<br />
The Extant Phylogenetic Bracket Approach<br />
The next question is, how do we obtain this requisite s<strong>of</strong>ttissue<br />
information? A method is outlined here and is presented<br />
in detail elsewhere (Witmer, 1992a, 1995a; see also Bryant and<br />
Russell, 1992, for an independently-devised but generally similar<br />
approach). The approach is firmly grounded in basic phylogenetic<br />
principles, somewhat resembling two-pass systems <strong>of</strong><br />
a posteriori character optimization (Wiley et al., 1991; Witmer,<br />
1995a). The only source <strong>of</strong> direct information about s<strong>of</strong>t tissues<br />
and their relationships to the skeleton are extant taxa, and thus<br />
they must be components <strong>of</strong> the analysis. The most relevant<br />
extant taxa are the two most-proximal living outgroups <strong>of</strong> the<br />
fossil taxon <strong>of</strong> interest (Fig. 3A). Figure 3B presents a topology<br />
in which there has been a rotation about the outgroup node such<br />
that the extant taxa now flank the fossil taxon. This rotation is<br />
simply a heuristic device that highlights a central objective <strong>of</strong><br />
a<br />
C<br />
2<br />
Extant Phylogenetic Bracket<br />
([I C,<br />
([I<br />
C<br />
C<br />
.-( w<br />
.rl .rl C<br />
C CI) CI) c<br />
([I<br />
([I<br />
C 0 CI)<br />
C<br />
m<br />
C<br />
2<br />
X 0 0 X<br />
a, b-4 4-r a,<br />
--3<br />
FIGURE 3. The Extant Phylogenetic Bracket Approach. A, phylogenetic<br />
relationships <strong>of</strong> a fossil taxon and its first two extant sister groups.<br />
B, rotation about the outgroup node in A brings the extant taxa to the<br />
periphery, forming the Extant Phylogenetic Bracket (EPB). C, cladogram<br />
showing the inference <strong>of</strong> s<strong>of</strong>t-tissue attributes in fossil taxa using<br />
the EPB approach. Similarities in s<strong>of</strong>t tissues and osteological correlates<br />
between the extant taxa are hypothesized as having been present in their<br />
common ancestor, the bracket ancestor (hatched arrows). If the fossil<br />
taxa possess the osteological (skeletal) correlates, then the hypothesis<br />
<strong>of</strong> homology survives the congruence test. If the s<strong>of</strong>t tissues and osteological<br />
correlates indeed are causally associated, then the s<strong>of</strong>t tissue<br />
can be inferred in the fossil taxa with confidence. (Modified from Witmer,<br />
1992a. 1995a.)<br />
bracket<br />
ancestor