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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

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