MESOZOIC MARINE REPTILE DISPARITY
reptile disparity was quantified using the PCOa scores of taxa from the first 11 ordination axes (Table 2), based on the sum of variances and the sum of ranges metrics. To derive a trajectory of disparity through time, we performed calculations on taxa binned in
variation. The scores (coordinates) of the taxa on PC axes 1, 2, and 3 were used to construct morphospaces because these axes contribute the most to overall absolute variance. Only the first 11 axes were retained for statistical tests and disparity calculations, because higher axes were associated with negligible amounts of absolute variance (<1%) or negative eigen- values (Table 2). To ensure that the presence of negative eigenvalues did not distort the ordinated dissimilarity among forms, we performed correlation tests on the computed intertaxon distances and ordination distances (Friedman 2012). Results show strong correlation between the distance measures (Pearson’s product-moment correlation: r=0.982, p<0.001), suggesting the PCOa accurately preserves most of the underlying distances between taxa without major distortion (Supplementary Fig. 1). Nonparametric multivariate analyses of variance (NPMANOVA) were carried out on the 11 retained PCOa axes to test for separation by temporal and group bins in morphospace (Ruta et al. 2013). The above-described procedureswere repeated separately for each of the five major clades to generate multivariate axes of variation for subsequent disparity calculations in each group (see below). Temporal Disparity Calculations.—Marine
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16 Mesozoic time intervals. The sum of ranges calculations were rarefied to account for differing sample sizes, using both the average sample size of all bins (n=17) and the smallest sample size (n=5). Statistical tests for differences in disparity between time bins are based on 95% confidence intervals, paired- sample t-tests based on multivariate variance (Anderson et al. 2011), and ratios of marginal likelihoods for variance (Finarelli and Flynn 2007). Confidence intervals associated with calculated disparity values were generated by bootstrapping with 10,000 replications. Ratios of marginal likelihoods were used exclusively to identify shifts in disparity between successive time intervals based on the procedure outlined in Finarelli and Flynn (2007) and Anderson et al. (2011). We assessed the statistical dependence between bin sample size and disparity by applying generalized differencing and assessing the strength and significance of correlations (e.g., Ruta et al. 2013). Partial disparity for each marine reptile group was also examined to see how they contribute to overall disparity through the Mesozoic (Foote 1993). All disparity calculations and significance tests were performed using the MDA Matlab® package (Navarro 2003) and in R (R Development Core Team 2011). Stratigraphic Binning.—For temporal disparity
TABLE 2. Eigenvalues and percentages of variance associated with each PCO axis from themultivariate analysis. The primary analysis is based on all 18 characters.
PC 1
2 3 4 5 6 7 8 9
10 11
Principal coordinate analysis Eigenvalue 2.6051
1.1496 1.0131
0.77706 0.32332 0.26086 0.15826 0.13675 0.13178 0.11304 0.094332
% variance 27.42
12.10 10.66 8.18 3.40 2.75 1.67 1.44 1.39 1.19 0.99
calculations and morphospace plots, taxa were binned in 16 Mesozoic time intervals. We used a combination of stage-level and composite bins, with ages in Myr: Olenekian (250–247.1), Anisian (247.1–241.5), Ladinian (241.5–237), Carnian (237–228.4), Norian (228.4–209.5), Hettangian– Sinemurian (201.3–190.8), Pliensbachian– Toarcian (190.8–174.1), Aalenian–Bathonian (174.1–166.1), Callovian–Oxfordian (166.1– 157.3), Kimmeridgian–Tithonian (157.3–145), Berriasian–Barremian (145–126.3), Aptian– Albian (126.3–100.5), Cenomanian–Turonian (100.5–89.8), Coniacian–Santonian (89.77–83.6), Campanian (83.6–72.1), and Maastrichtian (72.1–66). A Rhaetian bin was not included due to the lack of marine reptile fossils during this interval. Absolute ages for the stratigraphic stages are from Gradstein et al. (2012). The mean bin duration is 11Myr and the range is 22.9Myr.
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