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diff --git a/34233.txt b/34233.txt new file mode 100644 index 0000000..beaeab4 --- /dev/null +++ b/34233.txt @@ -0,0 +1,1157 @@ +The Project Gutenberg EBook of Cranial Osteiology of the Hylid Frog, +Smilisca baudini, by Linda Trueb + +This eBook is for the use of anyone anywhere at no cost and with +almost no restrictions whatsoever. You may copy it, give it away or +re-use it under the terms of the Project Gutenberg License included +with this eBook or online at www.gutenberg.org + + +Title: Cranial Osteiology of the Hylid Frog, Smilisca baudini + +Author: Linda Trueb + +Release Date: November 7, 2010 [EBook #34233] + +Language: English + +Character set encoding: ASCII + +*** START OF THIS PROJECT GUTENBERG EBOOK CRANIAL OSTEIOLOGY--SMILISCA BAUDINI *** + + + + +Produced by Chris Curnow, Tom Cosmas, Joseph Cooper and +the Online Distributed Proofreading Team at +https://www.pgdp.net + + + + + + + + + + ===================================================================== + UNIVERSITY OF KANSAS PUBLICATIONS + MUSEUM OF NATURAL HISTORY + + ------------------- + + Volume 18, No. 2, pp. 11-35 + + ------------------------- October 15, 1968 -------------------------- + + + Cranial Osteology of the Hylid Frog, + Smilisca baudini + + BY + + LINDA TRUEB + + + + UNIVERSITY OF KANSAS + LAWRENCE + 1968 + + + + + UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY + + Editors of this number: Frank B. Cross, William E. Duellman, + Philip S. Humphrey + + Volume 18, No. 2, pp. 11-35 + Published October 15, 1968 + + + UNIVERSITY OF KANSAS + Lawrence, Kansas + + + + PRINTED BY + ROBERT R. (BOB) SANDERS, STATE PRINTER + TOPEKA, KANSAS + 1968 + [Union Label] + 32-3686 + + + + + Cranial Osteology of the Hylid Frog, + Smilisca baudini + + BY + + LINDA TRUEB + + + + +INTRODUCTION + + +The paucity of descriptive cranial anatomical work dealing with hylid +frogs was pointed out by Trueb (1966) in her paper describing the +cranial osteology of _Hyla septentrionalis_. Comparative studies on +the cranial osteology of the genus _Smilisca_ (Duellman and Trueb, +1966), along with other more brief descriptions, reveal variation +among cranial characters of hylids. Since these external characters +have been useful in defining species, species groups, and genera, it +seems worthwhile to pursue correlated studies on internal cranial +structure. The following account dealing with the Neotropical tree +frog, _Smilisca baudini_ Dumeril and Bibron, 1841, is the first +published description of the internal cranial anatomy of a hylid frog, +and supplements the recent account (Duellman and Trueb, 1966) of +external cranial osteology of the same species. Comparative studies of +hylid skulls are expected to yield information of taxonomic +importance. + +I am grateful to Richard J. Baldauf of Texas A & M University and +William E. Duellman of the University of Kansas for critically reading +the manuscript and offering helpful suggestions. The findings reported +here result from research on Middle American hylids supported by a +grant from the National Science Foundation (GB-1441) to William E. +Duellman. + + +Materials and Methods + +The serial sections illustrated beyond are from an adult male of +_Smilisca baudini_ (KU 89924) having a snout-vent length of 53.0 mm. +and a head width (measured at angle of jaws) of 17.0 mm. The specimen +was collected 5.2 kilometers east-southeast of Cordoba, Veracruz, +Mexico. Transverse sections were cut at thicknesses of 10 and 15 +microns on a rotary microtome and stained according to the technique +described by Baldauf (1958). Cleared and stained specimens and dried +skeletons also were used. Figure 1 is based on KU 68183-4 and Fig. 9 +on KU 55614. All other drawings are made from KU 89924. In all +cross-sectional figures, bone is represented by solid black, cartilage +by stippling, and connective tissue by cross-hatching. Unless otherwise +noted all descriptions are given in an anterior-posterior sequence. + +Commonly accepted English terms are used. For example, dentary is used +in preference to dentale and maxillary process instead of processus +maxillaris. If no commonly accepted English term is available for a +given structure, the Latin name is retained. For example, the +cartilaginous plate separating the cavum principale from the cavum +medium is termed the lamina superior. + + [Illustration: FIG. 1. Partially disarticulated skull (left + frontoparietal and nasal removed) of _Smilisca baudini_, KU 68183, + [Female] x 4. Abbreviations: _al. proc._, alary process of + premaxillary; _ant. sq._, anterior arm of squamosal; _epi. em._, + epiotic eminence; _exocc._, exoccipital; _fpar._, frontoparietal; + _fpar. fon._, frontoparietal fontanelle; _max._, maxillary; _nas._, + nasal; _pal._, palatine; _pal. proc._, palatine process; _pasph._, + parasphenoid; _pmax._, premaxillary; _pvom._, prevomer; _post, + sq._, posterior arm of squamosal; _pro._, prootic; _pter._, + pterygoid; _qj._, quadratojugal; _spmax._, septomaxillary; _sept, + nas._, septum nasi; _spheth._, spnenethmoid; _vent, sq._, ventral + arm of squamosal.] + + + + +DESCRIPTION OF INTERNAL CRANIAL OSTEOLOGY + + +Olfactory Region + +_Alary cartilage._--The anterior end of the alary cartilage (_al. c._, +Figs. 2-5) lies within the posterior concavity of the alary process +(_al. proc._, Figs. 1-3) of the premaxillary (_pmax._). In posterior +sections the cartilage assumes a dorsolateral position (Fig. 3), +ventral and slightly lateral to the tectum nasi. The alary cartilage +remains narrowly separated from the tectum nasi but fuses +ventromedially with the septum nasi and forms a nearly complete +cartilaginous capsule around the anterior end of the cavum principale. +Posterior to the anterior end of the cavum medium and the lamina +superior, the alary cartilage separates ventrally from the lamina. In +subsequent posterior sections, the cartilage, arcuate in cross +section, becomes progressively smaller and terminates at the level of +the union of the medial and lateral recesses of the cavum inferior. + +_Prenasal cartilages._--The superior prenasal cartilage is small; it +lies adjacent to the posterodorsal surface of the alary process of the +premaxillary, and anterior to the alary cartilage. The inferior +prenasal cartilage (_inf. pnas. c._, Figs. 2-6, and 8) appears +posterior to the appearance of the alary cartilage. The anterior +terminus lies at the base of the alary process; the cartilage extends +dorsally (Fig. 3) along the posterior surface of the alary process and +then curves posterodorsally and joins the solum nasi medioventral to +the posterior end of the septomaxillary (Fig. 8d). + +_Tectum nasi._--The anterolateral corner of the tectum nasi (_tect. +nas._, Fig. 2) appears just posterior and dorsomedial to the anterior +end of the alary cartilage. The anterior process is short; it fuses +medially with the septum nasi forming a complete roof to the cavum +principale (Figs. 3 and 4). The oblique cartilage (_obl. c._) diverges +laterally from the tectum nasi just posterior to the terminus of the +alary cartilage (Fig. 6). Medially, the tectum nasi persists, overlaid +by the nasal bone laterally. + +_Septum nasi._--Posterior to the appearance of the septum nasi (_sept, +nas._) and its union with the tectum nasi (Fig. 3), the septum +abruptly expands across the width of the skull medial to the alary +cartilage. The septum is entirely cartilaginous posterior to the level +of the olfactory eminence, except for a small amount of secondary +membranous ossification dorsomedially at a level anterior to the nasal +bones. Perichondral ossification commences in the dorsal part of the +septum nasi at the level of the olfactory eminence. Endochondral +ossification first appears dorsally in the vertical part of the septum +at the level of the internal nares. Ossification of dorsal parts of +the septum precedes ossification of ventral parts. Perichondral +ossification of the ventral part of the septum nasi is first noted at +the level of transition between the planum antorbitale and solum +nasi. Perichondral ossification gradually gives way to endochondral +ossification posteriorly. + + [Illustration: FIGS. 2-5. Transverse sections through anterior end + of skull: 2) anterior level of inferior prenasal cartilage; 3) + anterior level of internasal septum; 4) olfactory capsule at + anterior level of cavum principale; 5) olfactory capsule at + anterior level of cavum inferior. Abbreviations: _al. c._, alary + cartilage; _al. proc._, alary process of premaxillary; _cav. med._, + cavum medium; _cav. prin._, cavum principale; _cr. int._, crista + intermedia; _inf. pnas. c._, inferior prenasal cartilage; _l. + inf._, lamina inferior; _l. sup._, lamina superior; _max._, + maxillary; _pmax._, premaxillary; _r. etx. n. f._, ramus externus + narium foramen; _r. med. n. f._, ramus medialis narium foramen; + _rec. lat._, recessus lateralis; _sept. nas._, septum nasi; _sol. + nas._, solum nasi; _spmx._, septomaxilla; _tect. nas._, + tectum nasi.] + + [Illustration: FIGS. 6-7. Transverse sections through olfactory + capsule: 6) posterior level of cavum medium; 7) anterior level of + prevomer. Abbreviations: _cav. inf._, cavum inferius; _cav. med._, + cavum medium; _cav. prin._, cavum principale; _cr. sub._, crista + subnasalis; _ext. nar._, external nares; _inf._, infundibulum; + _inf. pnas. c._, inferior prenasal cartilage; _l. inf._, lamina + inferior; _l. sup._, lamina superior; _max._, maxillary; _nas._, + nasal; _ncl. dt._, nasolacrimal duct; _obl. c._, oblique cartilage; + _p. fac._, pars facialis; _p. pal._, pars palatina; _pvom._, + prevomer; _rec. med._, recessus medialis; _sept. nas._, septum + nasi; _sol. nas._, solum nasi; _spmax._, septomaxillary; + _tect. nas._, tectum nasi.] + +_Nasal cavities and associated structures._--The cavum principale +(_cav. prin._, Fig. 4) is the most anterior of the nasal cavities. It +first appears within the capsule bordered dorsally by the tectum nasi, +medially and ventrally by the septum nasi, and laterally by the alary +cartilage. The cavity extends posteriorly within recesses of the +sphenethmoid to the level at which the septum nasi terminates. + +The cavum medium (_cav. med._, Fig. 5) lies ventral and slightly +posterior to the anterior end of the cavum principale. It appears +slightly anterior to the septomaxillary at the level of the foramen +ramus externus narium and ramus medialis narium. The appearance of the +cavum medium within the ventrolateral extension of the septum nasi +divides the latter into an upper component, the lamina superior +(_l. sup._) lying between the cavum principale and cavum medium, and a +lower part, the lamina inferior (_l. inf._) lying ventral to the cavum +medium. As the cavum medium increases in width in posterior sections, +the lamina superior and lamina inferior lose their lateral connection. +The lateral part of the cavum medium diverges in the region of the +external nares as the nasolacrimal duct (_ncl. dt._) and the cavum +medium becomes confluent with the cavum principale (Fig. 6). The +posterior end of the cavum medium lies at the level of the posterior +terminus of the septomaxillary. + +Slightly posterior to the anterior end of the cavum medium the foramen +for the ramus externus narium (_r. ext. n. f._) and ramus medialis +narium (_r. med. n. f._) opens ventromedially into the floor of the +septum nasi (Fig. 4). The ventral closure of the floor of the foramen +completes the solum nasi, marks the anterior end of the recessus +medialis of the cavum inferior, and differentiates the roof of the +recess, the crista intermedia (_cr. int._), from the solum (Fig. 5). +The crista intermedia joins the laminae superior and inferior and +joins them for a short distance to the septum nasi medially. The +anterolateral part of the cavum inferior (_cav. inf._) lies +ventrolateral to the cavum medium, and extends medially to join the +medial recess. The fusion of the two recesses of the cavum inferior +completely separates the lamina inferior from the solum nasi (Fig. 6). + +Near the level of the union of the recessus lateralis and recessus +medialis of the cavum inferior, the crista intermedia separates from +the septum nasi, and the lamina superior diverges at its mid-width to +accommodate the septomaxillary (_spmax._) (Figs. 5 and 6). The lateral +remnant of the lamina superior and transition zone between the lamina +superior and lamina inferior is short and is quickly replaced by +connective tissue. Slightly posterior, the distal edge of the lamina +inferior diverges laterally as a small process, which lies dorsal to +the pars facialis (_p. fac._) of the maxillary. This part of the +lamina inferior terminates posteriorly at the level of confluence +between the cavum principale and lateral recess of the cavum inferior. + +Slightly posterior to the divergence of the crista intermedia from the +septum nasi, the crista terminates, thereby separating the lamina +superior and the lamina inferior from one another (Figs. 6 and 8). The +lamina superior terminates at the level of the infundibulum (_inf._, +Fig. 7), whereas the lamina inferior extends posterolaterally, +increases greatly in depth and joins the laterally ascending oblique +cartilage (_obl. c._) to form the planum terminale (_pla. ter._, +Fig. 10). + +The crista subnasalis (_cr. sub._, Fig. 6) differentiates from the +lateral edge of solum nasi adjacent to the maxillary in sections just +posterior to the confluence of the recessus medialis and the recessus +lateralis. The crista persists as a rod of cartilage which gradually +diminishes in size and terminates at the posterior level of the +septomaxillary. + +_The septomaxillary._--The septomaxillary (_spmax._, Fig. 9a-c) is a +triradiate bone. The anterior terminus is a thin sliver of bone +oriented horizontally between the cavum principale and cavum medium +and lateral to the lamina superior (Figs. 5 and 8a). This anterior +ramus of the septomaxillary increases in size posteriorly and diverges +medially into a medial ramus (_med. r. spmax._) and lateral ramus +(_lat. r. spmax._, Fig. 9a-b) to accommodate the confluence of the +cavum principale and cavum medium (Figs. 6 and 8b-c). The small medial +ramus is associated with the distal end of the lamina superior whereas +the lateral ramus lies dorsal to the lateral margin of the cavum +medium. Just anterior to the anterior end of the nasolacrimal duct, +the ventral ramus of the septomaxillary (_vent. r. spmax._, Fig. 9c) +is present in cross-sections ventral to the cavum medium. The ventral +ramus joins the horizontal and dorsal rami of the septomaxillary at +the anterior end of the nasolacrimal duct. The medial branch terminates +posteriorly at the level at which the cavum principale joins the cavum +inferior. The lateral ramus of the septomaxillary terminates +posteriorly at the level at which the recessus medialis diverges from +the recessus lateralis posteriorly and cavum principale and recessus +lateralis are confluent. + +_Planum terminale._--Posterior to the infundibulum the lamina inferior +and oblique cartilage join to form the planum terminale (_pla. term._) +which lies lateral to the cavum principale (Fig. 10). The lamina +inferior diverges ventrally from the planum terminale anterior to the +olfactory eminence. The planum terminale is restricted ventrally and +terminates at the level of the olfactory eminence (_olf. em._, +Fig. 11). + + [Illustration: FIG. 8. Transverse sections through olfactory + capsule in region of septomaxillary: _a_) anterior terminus of + septomaxillary; _b_) medial divergence of septomaxillary; _c_) + dorsal ramus of septomaxillary; _d_) posterolateral terminus of + septomaxillary. Encircled numbers represent the nasal cavities as + follows: 1) cavum principale; 2) cavum medium; and 3) cavum + inferius. Abbreviations: _al. c._, alary cartilage; _ant. spmax._, + anterior end of septomaxillary; _cr. int._, crista intermedia; + _dor. r. spmax._, dorsal ramus of septomaxillary; _ext. nar._, + external nares; _inf. pnas. c._, inferior prenasal cartilage; _l. + inf._, lamina inferior; _l. sup._, lamina superior; _lat. r. + spmax._, lateral ramus of septomaxillary; _med. r. spmax._, medial + ramus of septomaxillary; _nas._, nasal; _nlc. dt._, nasolacrimal + duct; _obl. c._, oblique cartilage; _pvom._, prevomer; _sept. + nas._, septum nasi; _sol. nas._, solum nasi; _spmax._, + septomaxillary; _tect. nas._, tectum nasi.] + + [Illustration: FIG. 9. Septomaxillary drawn from cleared and + stained specimen of _Smilisca baudini_, KU 55614: _a_) dorsal; + _b_) ventral; _c_) lateral. In each example, the anterior end lies + to the left. Abbreviations: _dor. r. spmax._, dorsal ramus of + septomaxillary; _lat. r. spmax._, lateral ramus of septomaxillary; + _med. r. spmax._, medial ramus of septomaxillary; _vent. r. + spmax._, ventral ramus of septomaxillary.] + +_Anterior and posterior maxillary processes._--The anterior end of the +anterior maxillary process (_ant. max. proc._) lies within the +maxillary at the level of the posterior terminus of the planum +terminale. The anterior maxillary process diverges medially from the +maxillary (Fig. 13) and expands dorsally along the medial face of the +pars facialis to meet the planum antorbitale just anterior to the +transition zone between the latter and the solum nasi. Posterior to +the transition zone, the planum antorbitale disappears and the +posterior maxillary process is restricted ventrally along the pars +facialis of the maxillary. Posteriorly the cartilage is associated +with the pterygoid, where it is known as the pterygoid process +(_pter. proc._, Fig. 14). + +_Planum antorbitale._--The anterior terminus of the planum antorbitale +(_pla. ant._) lies medial to the ventrolateral part of the nasal and +lateral to the internal nares (Fig. 12). It abruptly expands dorsally +along the medial face of the nasal to join the tectum nasi +dorsolaterally; somewhat posteriorly the planum antorbitale joins the +anterior maxillary process ventrally at the posterior margin of the +internal nares. + + [Illustration: FIGS. 10-11. Transverse sections through posterior + part of olfactory capsule: 10) region of planum terminale; 11) + anterior region of olfactory eminence. Abbreviations: _cav. p._, + cavum principale; _max._, maxillary; _nas._, nasal; _nlc. dt._, + nasolacrimal duct; _olf. em._, olfactory eminence; _p. fac._, pars + facialis; _p. pal._, pars palatina; _pla. ter._, planum terminale; + _pvom._, prevomer; _rec. lat._, recessus lateralis; _sept. nas._, + septum nasi; _sol. nas._, solum nasi; _tect. nas._, tectum nasi.] + + [Illustration: FIG. 12. Transverse section through the olfactory + capsule in region of planum antorbitale. Abbreviations: + _cav. prin._, cavum principale; _int. nar._, internal nares; _max._, + maxillary; _nas._, nasal; _olf. em._, olfactory eminence; + _p. fac._, pars facialis; _p. pal._, pars palatina; _pal._, palatine; + _pla. ant._, planum antorbitale; _pvom._, prevomer; _sept. nas._, + septum nasi; _sol. nas._, solum nasi; _tect. nas._, tectum nasi.] + +_External dermal bones associated with the olfactory region._--The +association of the premaxillary (_pmax._) to the nasal cartilages is +described in preceding sections. The premaxillaries are separated from +each other medially and from the maxillaries laterally by dense +connective tissue. Anteriorly, the maxillary (_max._) bears a small +palatine process (_pal. proc._, Fig. 1) and a long, delicate pars +facialis (_p. fac._, Fig. 6), which terminates dorsally at the level +of the lamina inferior. Posterior to the transition zone between the +planum antorbitale and solum nasi the pars facialis is greatly +reduced. The pars palatina (_p. pal._, Fig. 6) persists to the +posterior part of the orbit. + +The anterior end of the prevomer (_pvom._, Fig. 1) is associated with +the venter of the solum nasi at the level of the infundibulum just +posterior to the incorporation of the inferior prenasal cartilage into +the solum (Fig. 7). The prevomer expands dorsally around the distal +end of the solum to provide a bony lateral support for the olfactory +eminence (Figs. 10 and 11). A distal wing of the prevomer forms the +bony anterior and medial margins of the internal nares. + +The palatine (_pal._, Figs. 1 and 12) lies in connective tissue medial +and adjacent to the pars facialis. At its maximum size the palatine +forms the bony posterior margin of the internal nares and extends +dorsomedially from the pars palatina to the distal part of the solum +nasi. + +The nasal (_nas._, Fig. 1) is a thin bone overlying the tectum nasi +anteriorly (Fig. 7). It expands laterally to form a complete roof over +the cavum principale (Fig. 10). In the region of the internal nares, +the nasal forms the lateral wall of the cavum principale (Fig. 12). + + +Sphenethmoid Region + +Posterior to the transition zone between the planum antorbitale and +solum nasi, the sphenethmoid (_spheth._, Fig. 1) is fully ossified +medially, the lateral parts of the bone at this level are only +ossified perichondrally. The septum nasi persists at the anterior +level of the orbit and terminates just anterior to the orbitonasal +foramen (_orbnas. f._) and the anterior end of the parasphenoid +(_pasph._, Fig. 13). The orbitonasal foramen is moderately large, has +a complete bony margin, and is located at the dorsolateral corner of +the braincase. + + [Illustration: FIG. 13. Transverse section through sphenethmoid + region at level of orbitonasal foramen. Abbreviations: _ant. max. + proc._, anterior maxillary process; _max._, maxillary; _orbnas. f._, + orbitonasal foramen; _pasph._, parasphenoid; _spheth._, sphenethmoid.] + +At the level of the orbitonasal foramen, the sphenethmoid is entirely +ossified except for a small dorsolateral extension. This distal +extension expands laterally in posterior sections as the braincase is +increased to its maximum width at the mid-length of the orbit; the +cartilaginous margin is retained throughout the length of the +sphenethmoid. + +The bony dorsomedial part of the sphenethmoid diverges, forming the +anterior border of the frontoparietal fontanelle (_fpar. fon._, Figs. +1 and 14). The entire fontanelle is covered with a layer of dense +connective tissue continuous with that in which marginal bones and +cartilage of the sphenethmoid lie, and which is discrete from the +lower dermal layer of the overlying skin. At this level the braincase +is U-shaped in cross-section. Ossification terminates first in the +ventrolateral corners, followed by the lateral and dorsolateral areas. +The bony support of the latter area is furnished by the lamina +perpendicularis (_lam. perp._) of the frontoparietal (_fpar._, Fig. +14). Cartilage appears in the ventral part of the sphenethmoid in +posteromost sections; at the posterior levels of the orbit the +sphenethmoid is entirely cartilaginous. + + +Orbital, Otic, and Occipital Regions + +_Orbital region._--The sclera (_scl._, Fig. 14) of the eye is +cartilaginous. The optic foramen (_opt. f._) is large and lies in +connective tissue at the posterior limits of the orbit and +sphenethmoid. At the posterior levels of the foramen the dorsolateral +cranial roof cartilages, taeniatecti marginales (_t. t. mar._) +converge medially to form the posterior margin of the frontoparietal +fontanelle and the tectum synoticum (_tect. syn._) of the occipital +region. At the posterior levels of the orbit the bursa angularis oris +(_b. ang. o._, Fig. 14) is present adjacent to the maxillary. + +_Nerve foramina of otic and occipital regions._--The trochlear foramen +lies within the bony margins of the optic foramen. The trochlear nerve +is located posterodorsal to the optic tract, and separated from the +latter by connective tissue. The oculomotor foramen (_ocul. f._) lies +in connective tissue posterior and ventral to the optic foramen (Fig. +15). Anteriorly, dorsally, and ventrally the foramen has a bony margin +formed by the prootic (_pro._); posteriorly, only a thin layer of +connective tissue separates the oculomotor from the large prootic +foramen (_pro. f._). The latter is bordered by bone dorsally and by +cartilage ventrally (Fig. 16). Posteriorly, bone separates the prootic +foramen from the anterior acoustic foramen (_ant. acus._ _f._), +through which the ramus acusticus anterior and medius pass (Fig. 17). +An extremely narrow bridge of cartilage separates the anterior +acoustic foramen from the larger posterior acoustic foramen (_post. +acus. f._). The latter has a bony posterior margin and is widely +separated from the bony jugular foramen (_jug. f._) posteriorly +(Fig. 19). + + [Illustration: FIGS. 14-15. Transverse sections through skull: 14) + at level of optic foramen; 15) at level of oculomotor foramen. + Abbreviations: _angspl._ angulosplenial; _ant. r. pter._, anterior + ramus of pterygoid; _ant. sq._, anterior arm of squamosal; _b. ang. + o._, bursa angularis oris; _fpar._, frontoparietal; _fpar. fon._, + frontoparietal fontanelle; _l. perp._, lamina perpendicularis of + frontoparietale; _max._, maxillary; _Mc. c._, Meckel's cartilage; + _ocul. f._, oculomotor foramen; _opt. f._, optic foramen; _pasph._, + parasphenoid; _psdbas. proc._, pseudobasal process; _pter. proc._, + pterygoid process; _scl._, sclera; _t. t. mar._, taenia tecti + marginalis; _tymp. r._, tympanic ring.] + +_Pterygoid._--The anterior terminus of the pterygoid (_pter._, Fig. 1) +appears at approximately the mid-length of the orbit as a small +arcuate bone closely applied to the posterior maxillary process. +Farther posteriorly the maxillary decreases in size, and the pterygoid +and posterior maxillary process diverge medially from it. Posterior to +this point of divergence, the posterior maxillary process is known as +the pterygoid process (_pter. proc._). The anterior terminus of the +quadratojugal (_qj._) lies medial to the maxillary at the level of the +oculomotor foramen (Fig. 14). + +_Otic region._--The anterior end of the otic capsule (_ot. cap._) is +present at the anterior level of the oculomotor foramen. The anterior +terminus of the pseudobasal process (_psdbas. proc._) lies within the +medial portion of the pterygoid at the posterior border of the +oculomotor foramen (Fig. 15). The pseudobasal process abruptly +increases in size. At the level of the prootic foramen (Fig. 16) the +medial branch of the pterygoid diverges from the posterior ramus and +is closely applied to the medial surface of the pseudobasal process. +The otic process extends along the medial surface of the squamosal +from the dorsolateral edge of the pseudobasal process, and then +expands medially to meet the bony edge of the otic capsule and form +the crista parotica. Posterior to the formation of the crista +parotica, the ventral part of the otic process splits. The medial part +forms the ventrolateral ledge of the otic capsule (_vl. l. ot. c._, +Fig. 20a-f), whereas the lateral part moves ventrad in association +with the ventral arm of the squamosal and fuses with the pterygoid +process posteriorly. + +Posterior to the bony closure of the prootic foramen, the ventromedial +part of the pseudobasal process joins the prootic and forms the +ventrolateral edge of the otic capsule. The posterior terminus of the +medial branch of the pterygoid lies ventral to the lateral part of the +otic capsule. The posterior branch of the otic process merges with the +pterygoid process ventrally. + + [Illustration: FIGS. 16-17. Transverse sections through otic + region: 16) at level of prootic foramen; 17) at level of anterior + acoustic foramen. Abbreviations: _angspl._, angulosplenial; _ant. + acus. f._, anterior acoustic foramen; _cr. par._, crista parotica; + _fpar._, frontoparietal; _max._, maxillary; _Mc. c._, Meckel's + cartilage; _ot. cap._, otic capsule; _pasph._, parasphenoid; + _pro._, prootic; _psdbas. proc._, pseudobasal process; _pter._, + pterygoid; _pter. proc._, pterygoid process; _qj._, quadratojugal; + _sq._, squamosal; _tect. syn._, tectum synoticum; _tymp. r._, + tympanic ring.] + + [Illustration: FIGS. 18-19. Transverse sections through otic capsule: + 18) at level of posterior acoustic foramen; 19) at level of jugular + foramen. Abbreviations: _angspl._, angulosplenial; _corn. prin._, + cornu principalis; _cr. par._, crista parotica; _exocc._, exoccipital; + _fpar._, frontoparietal; _jug. f._, jugular foramen; _max._, maxillary; + _Mc. c._, Meckel's cartilage; _ot. cap._, otic capsule; _p. ext. pl._, + pars externa plectri; _p. int. pl._, pars interna plectri; _p. med. + pl._, pars media plectri; _pasph._, parasphenoid; _post. acus. f._, + posterior acoustic foramen; _postlat. cr. par._, posterolateral edge + of crista parotica; _pro._, prootic; _pter._, pterygoid; _pter. proc._, + pterygoid process; _quad. proc._, quadrate process; _qj._ + quadratojugal; _sq._, squamosal; _tect. syn._, tectum synoticum; + _tymp. r._, tympanic ring.] + +At the level of the anterior acoustic foramen the cornu principalis of +the hyale (_corn. prin._) appears as a lateral ledge at the +ventrolateral corner of the otic capsule (Fig. 20a-b). The cornu +principalis diverges from the ledge at the level of the abbreviated +bridge between the anterior and posterior acoustic foramina. In +posterior sections the cornu lies medial to the squamosal-pterygoid +process-pterygoid complex (Fig. 18). The posterior terminus of the +cornu lies at a level with that of the posterior acoustic foramen. + +The pars externa plectri (_p. ext. pl._, Fig. 20a-b) is cartilaginous +and first appears dorsal to the ventral arm of the squamosal in +association with the tympanic membrane. The pars externa plectri +expands dorsomedially and is fused briefly to the crista parotica by +the pars ascendens plectri (_p. asc. pl._, Fig. 20b). The pars interna +plectri (_p. int. pl._, Fig. 20b-f) is cartilaginous and appears +medial to the pars media plectri and the ventrolateral ledge of the +otic capsule at the level of the anterior acoustic foramen. The pars +media plectri (_p. med. pl._, Fig. 20b-f), a cartilage and bone +element, appears proximally at the dorsolateral edge of the otic +capsule and distally, ventral to the squamosal at a level between the +anterior and posterior acoustic foramina. At the level of the +posterior acoustic foramina the pars media plectri is bony, greatly +expanded in size, and joined to the pars interna plectri medially. + +The operculum (_op._, Fig. 20d-h) is cartilaginous and lies medial to +the lateral edge of the otic capsule between the pars interna plectri +and pars media plectri. The anterior end of the operculum (Fig. 20d) +lies at a level corresponding to the posterior part of the posterior +acoustic foramen. Posteriorly the operculum increases in size, and the +pars interna plectri and pars media plectri are reduced (Fig. 20e-f). +At a level corresponding to the posterior border of the posterior +acoustic foramen the medial portion of the pars interna plectri +disappears and leaves a small lateral rod of cartilage surrounded on +all but the ventral side by the operculum (Fig. 20f). The operculum +expands medially to merge with the main part of the otic capsule +(Fig. 20g). The lateral edge of the operculum expands ventrally and +then dorsomedially to form a complete tube. Slightly more posteriorly +the cartilaginous lateral edge of the otic capsule, lateral to the +operculum, dissipates into connective tissue and finally disappears, +leaving the posterior end of the operculum as the most distal element +of the otic capsule (Fig. 20h). + + [Illustration: FIG. 20. Transverse sections through otic capsule: + _a_) level of anterior ledge of otic capsule; _b_) anterior level + of pars interna plectri and pars ascendens plectri; _c_) level of + pars media plectri; _d-f_) successive levels of operculum and pars + media plectri; _g-h_) posterior levels of operculum. Abbreviations: + _corn. prin._, cornu principalis; _cr. par._, crista parotica; + _op._, operculum; _p. asc. pl._, pars ascendens plectri; _p. ext. + pl._, pars externa plectri; _p. int. pl._, pars interna plectri; + _p. med. pl._, pars media plectri; _sq._, squamosal; _tymp. r._, + tympanic ring; _vl. l. ot. c._, ventrolateral ledge of otic + capsule.] + +_Ossification in otic and occipital regions._--The otic region of the +cranium is largely unossified. At the level of the optic foramen +(Fig. 14) the floor of the neurocranium is cartilaginous but completely +underlaid by the bony parasphenoid. The taenia tecti marginales and +the tectum synoticum are covered dorsally and laterally by the +frontoparietals. Perichondral ossification representing the prootic +bone occurs at the margin of the optic foramen and somewhat +posteriorly over part of the floor of the neurocranium. Perichondral +and endochondral ossification occurs in the sides of the neurocranium +ventral to the lamina perpendicularis. This ossification expands +laterally until it meets the crista parotica dorsolaterally and forms +the dorsal part of the prootic bone. The anteroventral edge of the +otic capsule remains cartilaginous. Posteriorly, at the level of the +anterior acoustic foramen, endochondral ossification is meager and +restricted to the dorsomedial parts of the otic capsule, plus a small +amount in the neurocranial floor; perichondral ossification is +restricted to the peripheral areas showing endochondral ossification. +Posteriorly, endochondral ossification is restricted in the dorsal +part of the otic capsule but somewhat increased in the floor of the +capsule. The lateral part of the otic capsule posterior to the +terminus of the operculum and the ventromedial and dorsomedial parts +of the neurocranium remain unossified. + + +Articular Region + +In the anterior sections (at the level of the oculomotor foramen) the +angulosplenial (_angspl._) is a moderate-sized bone (Fig. 15). +Meckel's cartilage (_Mc. c._) is present as a small ovoid cartilage +lying dorsolateral to the angulosplenial. Posteriorly, Meckel's +cartilage is dorsal to the angulosplenial. The cartilage increases in +size at the level of the posterior acoustic foramen, and the +angulosplenial decreases in size posteriorly. At the level of the +posterior border of the posterior acoustic foramen, the maxillary +terminates and is replaced by the quadratojugal. The quadratojugal, +ventral arm of the squamosal, pterygoid process, pterygoid, and +Meckel's cartilage converge. At the level of the jugular foramen +(_jug. f._) (Fig. 19) the quadratojugal is incorporated into the +squamosal-pterygoid process-pterygoid complex. The complex is narrowly +separated by connective tissue from Meckel's cartilage ventrally. The +quadrate process (_quad. proc._) is represented by the cartilage +bordered dorsally by the pterygoid process and the ventral arm of the +squamosal, and ventrally by Meckel's cartilage. At the posterior +terminus of the skull all bony elements of the articular region +terminate, except for a small terminal part of the angulosplenial +underlying Meckel's cartilage. + + + + +SUMMARY + + +Since no accounts comparable to the preceding for _Smilisca baudini_ +are available for other hylid frogs, it is meaningless to attempt any +discussion dealing with character significance or variation within the +Hylidae. There is considerable literature treating bufonids, +leptodactylids, ranids, and various Old World genera (see Baldauf, +1955, for a review of these works). Likewise, a comparison at the +familial level based on the study of a single species seems inadequate +and premature. By way of summary and synoptic description a list of +cranial osteological characters of _Smilisca baudini_ is presented. +The items selected enable comparison with similar compilations by +other workers, and are based in part on my unpublished observations of +other hylids. + + 1. Compared to hylids not having integumentary-cranial + co-ossification, the dermal roofing bones of _Smilisca baudini_ + are extensive, and the skull is well-ossified internally. In + contrast to most casque-headed hylids (those having + integumentary-cranial co-ossification), the dermal roofing bones + are much less extensive, the dermal sphenethmoid (see Trueb, 1966, + p. 563) is absent, and internal ossification is less extensive. + + 2. The solum nasi is not ossified; the septum nasi is ossified only + posteriorly, and the olfactory eminence is supported by the + cartilaginous solum nasi and the bony prevomer. + + 3. The lingual process is absent. There is no palatal cartilage + isolated between the premaxillaries. + + 4. The anterior end of the cavum medium lies anterior to the cavum + inferius. + + 5. The septomaxillary is basically a U-shaped structure and has a + dorsal, anteriorly curved, ramus on the lateral branch and a + longitudinal loop of bone ventrally. + + 6. A distinct pars nasalis is absent on the maxillary. + + 7. A cartilaginous sclera is present. + + 8. The taenia tecta marginalis and the tectum synoticum are the only + roofing cartilages present. + + 9. The external part of the plectral apparatus (columella) is + directed anterolaterally. The pars ascendens plectri is fused + with the crista parotica. + + 10. The pseudobasal process is fused to the otic capsule. + + 11. The cornu principalis of the hyale fuses with the pseudobasal + process. + + 12. Two acoustic foramina are present. + + 13. The sphenethmoid and prootic are synchondrotically united. + + 14. The frontoparietal is separate from the prootic and exoccipital. + + 15. The prootic and exoccipital are fused. + + 16. A bursa angularis oris is present. + + + + +LITERATURE CITED + + + BALDAUF, R. J. + 1955. Contributions to the cranial morphology of _Bufo w. woodhousei_ + Girard. Texas Jour. Sci., 7(3):275-311. + + 1958. A procedure for the staining and sectioning of the heads of + adult anurans. Texas Jour. Sci., 10(4):448-451. + + DUELLMAN, W. E. AND L. TRUEB + 1966. Neotropical hylid frogs, genus Smilisca. Univ. Kansas Publ., + Mus. Nat. Hist., 17:281-375, pls. 1-12. + + TRUEB, L. + 1966. Morphology and development of the skull of the frog _Hyla + septentrionalis_. Copeia, 3:562-573. + + + +_Transmitted April 18, 1968._ + + + + * * * * * + +Transcriber's Notes + +Except for several minor typographical corrections which were made +(missing periods, commas, etc.) that are not detailed here, the text +presented is that which appeared in the original printed version. 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