The following description of development of the skull of Smilisca baudini is based on the examination of 12 cleared and stained specimens. In table 3 the cranial bones are listed in the left hand column in the approximate order of their appearance in the young frogs. Across the top of the table selected specimens designated by developmental stage or snout-vent length are listed. It should be noted that although each individual, from left to right, has an increasing number of ossified bones, the correlation with increasing size is imperfect; the precise ages of the individuals are unknown.

The first bones to appear are the septomaxillaries, frontoparietals, part of the exoccipital, and the parasphenoid in developmental stage 40. The frontoparietals are represented by two slender ossifications dorsomedial to the orbits; the septomaxillaries are present as small ossifications anterior to the nasal capsules (Pl. 1A). The parasphenoid is present as a faint median ossification, and the exoccipital shows some ossification.

Table 3.—The Order of Occurrence of Cranial Ossifications in the Skull of Smilisca baudini. Where Numbers Are Divided by a Slash Mark, the Left and Right Symbols Correspond to the Left and Right Sides of the Skull, Respectively.
BoneStage 40Stage 4412.6 mm.13.9 mm.32.0 mm.27.0 mm.20.1 mm.
FrontoparietalXXXXXXX
ParasphenoidXXXXXXX
SeptomaxillariesXXXXXXX
ExoccipitalsXXXXXXX
Squamosals—XXXXXX
Premaxillaries—XXXXXX
Maxillaries—XXXXXX
Nasals——XXXXX
Pterygoids——XXXXX
Vomers———XXXX
Palatines———XXXX
Quadratojugals———XXXX
Ethmoid————XXX
Columellas————XXX
Supraorbital Flanges—————XX
Proötics——————X
Vomerine Teeth——1/14/35/53/35/4
Maxillary Teeth—0/73/56/530/3130/2637/36
Premaxillary Teeth—2/43/35/57/68/68/7

The dentigerous bones are among the most rapidly developed, although not the first to appear. They are present in developmental stage 44 before metamorphosis is completed. The maxillaries bear a few teeth anteriorly and are ossified posteriorly to a point one-third of the distance from the anterior to the posterior edge of the orbit. Ossification lengthens the posterior termini of the maxillaries to the posterior edge of the orbit. In front of the anterior margin of the orbit, bone is proliferated dorsal to the main axes of the maxillaries and forms moderate dorsal maxillary flanges. The premaxillaries appear simultaneously with the maxillaries. Initially they are widely separated medially from each other, and laterally from the developing maxillaries; each bears two or three teeth, large dorsally blunt alary processes, and small palatine processes. The median and lateral edges of the prenasal processes lengthen heterochronously, causing the median edges to be longest and to lie slightly dorsal to the level of the septomaxillaries. After the maxillaries and premaxillaries develop, the vomers appear as small horizontal ossifications anterior to the parasphenoid. Ossification begins in the lateral flanges, then in the prevomerine processes, and lastly in the posterior dentigerous parts of the bones; the prevomerine processes are the last parts of the vomers to ossify completely.

Initially the frontoparietals are present as thin rods of ossification dorsomedial to the orbits; the frontoparietals extend from the anterior to the posterior end of the orbit by developmental stage 44. The anterior ends of the bones remain thin and pointed; ossification progresses medially from the midpoint of the length of the orbit and posteriorly to the level of the exoccipital; a median center of ossification joins the frontoparietals posteriorly, thereby forming the posterior border of the frontoparietal fontanelle. The supraorbital flanges of the frontoparietals do not appear until all other cranial bones are ossified, or nearly so. The most rapid ossification begins laterally at the posterior edge of the orbit and decreases anteriorly over the posterior half of the orbit. This differential rate of proliferation of bone results in the pattern of development of the supraorbital flanges shown in figure 7. The nasals appear as thin slivers of bone half way between the anterior ends of the frontoparietals and the end of the snout. As ossification proceeds the nasals assume a triangular shape in dorsal view. The anterior ends are pointed; the lateral margins are parallel to the maxillaries. The posteromedial points do not reach the lateral margins of the ethmoid, and the maxillary processes extend about three-fourths the distance from the bodies of the nasals to the maxillaries. Following the union of the frontoparietals posteriorly, the nasals widen anteriorly and are narrower at the midpoints of their long axes than anteriorly or posteriorly. With further ossification the maxillary processes extend to the maxillaries and form complete bony anterior margins to the orbits; the mid-parts of the nasals widen (Pl. 1B).

Fig. 7. Developmental sequence of the frontoparietal fontanelle and associated bony elements in Smilisca baudird: (A) KU 60026, ×5; (B) KU 85438, ×4; (C) KU 26328, ×3; (D) KU 68184, ×2.3.

The parasphenoid is the first of the palatal bones to appear. At metamorphosis the bone is well developed; the anterior tip is situated just in front of the anterior edge of the orbit, and posteriorly the lateral processes extend laterally beyond the ossified parts of the auditory region. The pterygoids do not appear until metamorphosis, when ossification is evident in only the mid-parts of the posterolateral arms. Ossification follows in the mid-parts of the anterolateral arms and occurs last in the pterygoid pedicles. The palatines do not appear until all three arms of the pterygoids are at least partly ossified. Ossification proceeds rapidly from the maxillaries medially to the unossified ethmoid, which is the last of the cranial bones to appear. Initially it is extremely shallow; dorsally it is widely separated from the nasals, and ventrally the posterior margin meets the anterior point of the parasphenoid. In dorsal view, ossification proceeds anteriorly between the nasals and posteriorly, ventral to the frontoparietals; ventrally, ossification proceeds posteriorly dorsal to the parasphenoid.