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The diverse selective pressures proposed to drive extreme derivations in the skull include specializations in feeding biology (1), habitat use (2), and locomotion (3). Sexual selection also is thought to influence head morphology because the skull often is sexually dimorphic in size and shape (4, 5).

The nonadaptive mechanisms of architectural constraint (i. The diversification of the skull usually results from changes in size or shape of preexisting elements or Intal Nebulizer Solution (Cromolyn Sodium Inhalation Solution)- FDA loss of bones (10), but the origin of novel structures also may be responsible for shifts in morphology (11).

In its most rudimentary form, additional membrane bone is deposited on the skeleton to form ridges and crests that produce a reticulate or pitting pattern on the surface of bones (exostosis), but, in extreme cases, hyperossification can lead to the formation of helmet-like protuberances (casquing) or coossification between the skeleton and dermis (14).

Hyperossification is thought to result from the heterochronic process of peramorphosis, which is the extension or acceleration of ancestral ontogenetic trajectories (15).

Frog skulls may be understudied because it has been assumed that the highly derived Bauplan and skeletal morphology of this clade are tightly conserved (19). Cranial hyperossification is expressed on five dermal elements (frontoparietal, nasal, premaxilla, maxilla, squamosal) and three novel bones (prenasal, internasal, dermal sphenethmoid) in frogs (29).

Alternately, hyperossification may arise as a byproduct of miniaturization (31). Using the most recent species-rich phylogeny of extant amphibian species (32) and extensive taxonomic sampling via high-resolution X-ray microcomputed tomography (158 species representing porno masturbation 54 described anuran families), we 1) evaluated the broad-scale patterns of skull shape diversity across all major frog lineages, 2) reconstructed the evolutionary Intal Nebulizer Solution (Cromolyn Sodium Inhalation Solution)- FDA of skull hyperossification, and 3) tested the ways in which body size, feeding biology, microhabitat use, and phragmotic defense behavior are associated with skull shape and interact with hyperossification.

Our results demonstrate that, although many lineages share a conserved skull shape, several highly divergent skull architectures have evolved repeatedly throughout the evolutionary history of frogs. Hyperossification has evolved independently many times and often cooccurs with divergent skull shapes.

Body size, microhabitat use, and feeding biology are each correlated with variation in head shapes, and vertebrate predation and phragmotic defense behavior cooccur with hyperossification and extreme skull shapes. We used three-dimensional (3D) geometric morphometric analyses on 36 fixed landmarks (SI Appendix, Fig. S1) in the R package geomorph version 3. S2), indicating that highly divergent shapes have evolved repeatedly.

Phylogenetic tree of frogs depicting the evolution of skull shape and hyperossification. Branch color gradient corresponds to maximum likelihood ancestral states of skull shape (PC2): Lineages with positive scores (white) have a narrow and flat skull, whereas lineages with negative scores (black) have a wide and tall skull. The size of each node point represents the posterior probability of the most probable ancestral state. Tip point colors correspond to hyperossification, diet, and microhabitat states for all 158 species.

A horizontal bar on the do you plan to quit drinking point indicates the presence of odontoid fangs, and a vertical bar on the microhabitat point indicates the presence of phragmotic behavior. Tip numbers 1 to 30 correspond Intal Nebulizer Solution (Cromolyn Sodium Inhalation Solution)- FDA species depicted in Fig.

Species tip labels are provided in SI Appendix, Fig. S4, and corresponding trait data are provided in Dataset S1. Phylomorphospace plots of (A and C) PC1 and PC2 and (B and D) PC2 and PC3 axes of shape variation exhibiting the diversity of skull morphology in frogs.

Points are colored by (A and B) microhabitat and (C and D) diet states. Point numbers 1 to 30 correspond to species in Fig. Ordinary least-squares regression lines are displayed for hyperossified species (black points, black line) and nonhyperossified species (white points, gray dashed line) to demonstrate the lack of slope differences between these two groups (see diversity in skull shape in results).

Species labels for all points are provided in SI Appendix, Fig. Hyperossification occurs in 44 of the Intal Nebulizer Solution (Cromolyn Sodium Inhalation Solution)- FDA anuran taxa in our dataset and is distributed across 39 Intal Nebulizer Solution (Cromolyn Sodium Inhalation Solution)- FDA and 17 families (Fig.

We used reversible-jump Markov chain Monte Carlo (MCMC) in RevBayes (34) to sample all five Markov models of phenotypic character evolution in proportion to their posterior probability, and the maximum a posteriori model of hyperossification evolution was the one-rate model with a posterior probability of 0. The model-averaged maximum a posteriori fd c yellow 5 state of frogs was nonhyperossified with a posterior johnson motor of 0.

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