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406 REPRODUCTIVE BIOLOGY AND EMBRYOLOGY OF CROCODILIANS<br />

Palate. The secondary palatal shelves are one-fourth closed at <strong>the</strong> beginning<br />

<strong>of</strong> Stage 18 <strong>and</strong> three-fourths closed at <strong>the</strong> end (Figs. 25B <strong>and</strong> C).<br />

This stage can be accurately subdivided by specifying <strong>the</strong> extent <strong>of</strong> secondary<br />

palate closure (Figs. 25B <strong>and</strong> C, <strong>and</strong> 28C-E). The upper jaw margin is<br />

straighter <strong>and</strong> less hooked than previous.<br />

Eye. The margins <strong>of</strong> <strong>the</strong> upper eyelid anlage extend over <strong>the</strong> superior<br />

rim <strong>of</strong> <strong>the</strong> iris forming a distinct groove between <strong>the</strong> eyelids <strong>and</strong> <strong>the</strong> eye,<br />

into which small instruments can be passed.<br />

Scales, Scutes, etc. Dorsal scalation is now marked.<br />

PericardiaI Sac. The bulge <strong>of</strong> <strong>the</strong> transparent pericardial sac is starting to<br />

be submerged into <strong>the</strong> ventral thoracic wall.<br />

Stage 19 (Figs. 20, 23, 250, 26B, 34E <strong>and</strong> F]<br />

Eye. Upper <strong>and</strong> lower eyelids are distinct. The anterior nictitating<br />

membrane anlage is discernible in <strong>the</strong> anterior corner <strong>of</strong> <strong>the</strong> eye (Fig. 26B).<br />

Caruncle. Two elevations <strong>of</strong> <strong>the</strong> caruncle have approximated each o<strong>the</strong>r<br />

at <strong>the</strong> tip <strong>of</strong> <strong>the</strong> snout (Fig. 26B), but <strong>the</strong> tissue between <strong>the</strong>m is thin,<br />

appearing transparent under incident illumination.<br />

Lower Jaw. The lower jaw lies behind <strong>the</strong> anterior margin <strong>of</strong> <strong>the</strong> upper<br />

jaw. Consequently, if <strong>the</strong> premaxillary bulges are large, <strong>the</strong> mouth opens<br />

(as in Fig. 24H). The tongue <strong>and</strong> floor <strong>of</strong> <strong>the</strong> mouth contents sag beneath<br />

<strong>the</strong> margins <strong>of</strong> <strong>the</strong> lower jaw.<br />

Limbs. Interdigital clefting has commenced producing slight marginal<br />

notches particularly in <strong>the</strong> foot plates (Fig. 23).<br />

Palate. The palate is almost completely closed (Fig. 250).<br />

Coloration. White flecks, representing underlying ossifications, are obvious<br />

on <strong>the</strong> margins <strong>of</strong> <strong>the</strong> upper <strong>and</strong> lower jaws <strong>and</strong> around <strong>the</strong> ears.<br />

External Genitalia. The end <strong>of</strong> <strong>the</strong> external genitalia has developed a<br />

globular swelling.<br />

Denticles. There are eight to nine denticles visible on each side <strong>of</strong> <strong>the</strong><br />

lower jaw. Henceforth, many m<strong>and</strong>ibular denticles appear <strong>and</strong> disappear<br />

so that <strong>the</strong>ir numbers are too variable to be used in staging.<br />

Species Differences. In C. johnsoni <strong>and</strong> C. porosus, <strong>the</strong> nostrils are elevated<br />

on a nasal disk. The lateral jaw margins have distinct notches where<br />

<strong>the</strong> primary <strong>and</strong> secondary palates closed (as in Figs. 24H <strong>and</strong> I); <strong>the</strong>se later<br />

accommodate <strong>the</strong> large fourth dentary teeth.<br />

Stage 20 (Figs. 20, 23, 24H, I, 25E, <strong>and</strong> 26C]<br />

Limbs. Nail anlagen develop rapidly in a specific sequence early in this<br />

stage (Fig. 23). They appear first on <strong>the</strong> most medial digit <strong>of</strong> <strong>the</strong> foot, <strong>the</strong>n<br />

on <strong>the</strong> neighboring two digits, <strong>the</strong>n on <strong>the</strong> most medial digit <strong>of</strong> <strong>the</strong> h<strong>and</strong><br />

<strong>and</strong> finally on <strong>the</strong> neighboring two medial h<strong>and</strong> digits. Consequently, nail<br />

anlagen are present on <strong>the</strong> most medial three digits <strong>of</strong> both <strong>the</strong> h<strong>and</strong>s <strong>and</strong><br />

STAGES OF EMBRYONIC DEVELOPMENT [AFTER EGG LAYING)<br />

407<br />

feet, despite <strong>the</strong> fact that <strong>the</strong> total number <strong>of</strong> digits varies between <strong>the</strong> two<br />

(Fig. 21). Interdigital clefting now extends along approximately one-fourth<br />

<strong>the</strong> length <strong>of</strong> <strong>the</strong> digits (Fig. 23). The outer two digits <strong>of</strong> <strong>the</strong> h<strong>and</strong> <strong>and</strong> <strong>the</strong><br />

outer digit <strong>of</strong> <strong>the</strong> foot never develop nails.<br />

Caruncle. The caruncle is now a solid structure due to consolidation <strong>of</strong><br />

<strong>the</strong> region between <strong>the</strong> two initial swellings (Fig. 26C).<br />

Lower Jaw. The lower jaw is in its adult relationship with <strong>the</strong> upper jaw<br />

(Figs. 24H <strong>and</strong> I).<br />

External Genitalia. The external genital primordium is now pointed<br />

with a distinct elevation <strong>of</strong> its tip.<br />

Palate. The palate is completely closed but <strong>the</strong> basihyal valve is not yet<br />

present (Fig. 25E).<br />

PericardiaI Sac. The pericardial sac is now one-fourth withdrawn into<br />

<strong>the</strong> ventral body cavity.<br />

Coloration. White flecks <strong>of</strong> ossification are present along <strong>the</strong> margins <strong>of</strong><br />

<strong>the</strong> upper <strong>and</strong> lower jaws, around <strong>the</strong> external auditory meatus, <strong>and</strong> in <strong>the</strong><br />

proximal <strong>and</strong> distal elements <strong>of</strong> <strong>the</strong> limbs.<br />

Scales <strong>and</strong> Scutes. Scale formation is marked dorsally <strong>and</strong> scutes are<br />

beginning to appear in <strong>the</strong> neck region behind <strong>the</strong> skull.<br />

Stage 21 (Figs. 20, 23, 25F, 260]<br />

Limbs. Interdigital clefting extends three-fourths <strong>of</strong> <strong>the</strong> way along <strong>the</strong><br />

digits. Phalanges can be distinguished in <strong>the</strong> digits (Fig. 23).<br />

Scales <strong>and</strong> Scutes. Scales are now visible on <strong>the</strong> ventral body wall as well<br />

as dorsally on <strong>the</strong> snout, neck, body, <strong>and</strong> tail. The dorsal neck scutes are<br />

clearly defined.<br />

Caruncle. The caruncle is a solid mass on <strong>the</strong> snout tip, but <strong>the</strong> tissue<br />

around <strong>the</strong> base <strong>of</strong> <strong>the</strong> caruncle is not differentiated from <strong>the</strong> o<strong>the</strong>r snout<br />

scales (Fig. 260).<br />

Palate. The superior basihyal valve flap is present at <strong>the</strong> posterior margin<br />

<strong>of</strong> <strong>the</strong> palate (<strong>and</strong> <strong>the</strong> inferior flap at <strong>the</strong> base <strong>of</strong> <strong>the</strong> tongue) <strong>and</strong> a<br />

plexus <strong>of</strong> palatal blood vessels is conspicuous (Fig. 25F).<br />

Pericardial Sac. The pericardial sac is one-half withdrawn into <strong>the</strong> body<br />

cavity.<br />

External Nares. Elevations for <strong>the</strong> constrictor nares muscles are evident.<br />

Eye. A white ring in <strong>the</strong> iris surrounds <strong>the</strong> outline <strong>of</strong> <strong>the</strong> lens <strong>of</strong> <strong>the</strong> eye<br />

<strong>and</strong> is overlapped by both upper a.nd lower eyelids.<br />

Stage 22 (Figs. 21, 23, 251]<br />

Coloration. Pigmentation is first visible on <strong>the</strong> margins <strong>of</strong> <strong>the</strong> upper<br />

jaw, along <strong>the</strong> ventral aspect <strong>of</strong> <strong>the</strong> flank, <strong>and</strong> on <strong>the</strong> proximal <strong>and</strong> distal<br />

elements <strong>of</strong> <strong>the</strong> limbs, but <strong>the</strong>re is little or no dorsal pigmentation.

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