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Caecilian Morphology

Caecilians are adapted primarily for burrowing and exhibit many of the same morphological characteristics shown by other fossorial ectotherms such as snakes, lizards, amphisbaenians, and salamanders. The Typhlonectids also show secondary adaptations for an aquatic or semi-aquatic existence. This page will cover some of the basics of caecilian morphology.

Body Form

Elongation of the body and the reduction or loss of limbs is a common adaptation in all burrowing tetrapods and caecilians are extreme examples of this. The result is a worm-like or snake-like appearance. Some aquatic species (Typhlonectes, Atretochoana, and Potomotyphlus) have a small, raised "fin" along the posterior portion of the body to aid in swimming and the result is an eel-like appearance. There is no trace of limbs or pelvic or pectoral girdles left in caecilians. By comparison, some caecilian-like salamanders still have at least two legs and some snakes and legless lizards still have remnants of legs under the skin. The fossil caecilian, Eocaecilia, had reduced legs.

The body is arranged in rings, called annuli (Figure 1). These annuli give caecilians their earthworm-like appearance. Additionally, the annuli may be further divided into secondary and tertiary annuli. Having secondary or tertiary annuli is considered to be a more primitive trait among caecilian families.

Although it can be difficult to conceptualize unless you have examined a caecilian in person, the tail makes up only a very small part of the body when it exists at all. In the majority of species, the body simply ends abruptly in a blunt terminal "shield" or "terminus" (Figure 2). Only the more primitive families of caecilians still retain a tail.

Body size varies markedly within the Gymnophiona, but is certainly constrained by a fossorial lifestyle. The smallest known caecilians are probably Idiocranium russeli from Cameroon with a largest recorded length of 14.4cm, though females as small as 9cm have been found brooding eggs. Individuals as small as 5.1cm have also been found with no trace of larval characteristics. On the Seychelles Archipelago, Grandisonia brevis (11.2cm) and G. dimunitiva (9.5cm) often are shorter in length than Idiocranium, but considerably thicker and heavier in body form.

The longest caecilian known is Caecilia thompsoni at 151.5cm but other species such as Caecilia tentaculata and Caecilia nigricans, each reaching lengths of 100cm or more, are much stouter and heavier.

The Skull

Caecilian skulls are strongly built and heavily ossified. The head acts as a battering ram as caecilians push and prod their way through soil. As a result, the skull is wedge- like and bones are heavily fused. Aside from sensory openings, the skulls of most caecilians are completely roofed with thick bone, a condition called stegokrotaphy (Figure 3). Other more primitive species still retain a partially open temporal region, a condition known as zygokrotaphy (Figure 4). A reduction of the number of bones in the skull is considered an advanced trait in caecilian families.

The Eyes

All caecilians have eyes, but they may be so reduced and covered by skin or bone that you may not be able to see them. Leading a fossorial life, the eyes have degenerated to various degrees in different species. Some species, such as Ichthyophis, have superficially placed eyes while other species such as Herpele (Figure 5) and Gegeneophis have eyes beneath the bones of the skull and have even had their eye sockets replaced by bone (Figure 6).

Comparative morphological studies suggest that there is a trend towards increased covering of the eye by skin or bone in conjunction with a loss of the lens and modifications of the retina. However, the retina and optic nerve remain intact and so it is likely that most caecilian eyes are still capable of photoreception.

Some caecilian keepers report that their animals do not show any reaction to lighting. Others do seem to notice an ability to detect light and report that their caecilians will often hide or jerk away when a bright light is suddenly turned on in their enclosure or is pointed at their eyes. They do not seem capable of detecting movement visually.

The Tentacles

A small sensory tentacle is always present on either side of the head between the eye and the nostril (Figure 7) and the exact positioning of the tentacles is highly variable among species. In most species, it can be protruded through an aperture in the skull while in others it is not protrusible (Typhlonectids). The family Scolecomorphidae is somewhat famous for having the tentacle in close association with the eye; when the tentacle is protruded, the eye is carried along outside the cranium on the tip of the tentacle.

The tentacle is a complex structure of many forms of tissues including nerves, muscles, ducts, and glands and is thought to function in chemoreception.

The Mouth and Teeth

In most caecilians, the mouth is not positioned directly at the end of the head (terminal), but rather is positioned slightly below (subterminal). This arrangement is also called "countersunk  and is considered an adaptation for burrowing (Figure 8). The most primitive caecilians still possess terminal mouths.

The upper and lower jaws each carry two rows of teeth, an inner row and an outer row. These teeth vary in size and shape depending on species and tooth counts change with age. Captive Typhlonectes have been known to mouth the hands and fingers of their keepers at feeding time, which is quite harmless and painless.

 

Nuchal Collars and Grooves

Immediately behind the head are two anatomical structures called nuchal collars, which are differentiated from each other and the rest of the body by nuchal grooves (figure 9). Somewhat similar in appearance to the clitellum of earthworms, the nuchal collars usually span 3 to 4 vertebrae and are most noticeable when viewing the caecilian from the side or below. The first nuchal groove marks the posterior border of the skull and the second marks the division between the two nuchal collars. The third nuchal groove marks the boundary between the second nuchal collar and the rest of the body. The nuchal collars may sometimes be difficult to distinguish in some species due to additional dermal folds along the dorsal surface.

The gills of larval caecilians often occur on the same plane as the second or third nuchal groove.

Internal Organs

Like snakes, amphisbaenians, salamanders, and legless lizards, the internal organs of caecilians have become greatly elongated. One or more of the lungs may be reduced or absent entirely, usually the left lung (Atretochoana eiselti lacks lungs entirely).

Jaw Muscles

All terrestrial vertebrates, except caecilians, have a single set of jaw-closing muscles. Caecilians have two sets of muscles (mandibular adductors and interhyoideus muscles) for this purpose and this is thought to be an adaptation for holding the jaws firmly closed while burrowing.

The Skin

Like other amphibians, caecilians do possess poison glands in the skin although the potency of such poisons is not yet well known. Toxic skin secretions have been observed for Typhlonectes compressicauda and hemolytic and cardiotoxic properties have been found in the skin secretions of Siphonops paulensis .

Unlike other extant amphibians, caecilians possess scales under the skin, which are composed of collagenous fibers covered with mineralized nodules. These can be found in the folds and grooves of the skin and usually increase in number posteriorly. Additionally, Caecilia sp. have a second type of scale which is embedded into the subdermal connective tissue.


References:

Cogger, H.G., & R.G. Zweifel. 2003. Encyclopedia of Reptiles and
     Amphibians: A Comprehensive Illustrated Guide by International Experts. Fog City
     Press, San Francisco.

Measey, J.G., Gower, D.J., Oomen, V.O., & M. Wilkinson. 2004. A Subterranean
     Generalist Predator: Diet of the Soil Dwelling Caecilian Gegeneophis ramaswamii
    
(Amphibia; Gymnophiona, Caecilidae) in Southern India. C.R. Biologies 327: 65-76.   

Pillai, R.S. & M.S. Ravichandran. 1999. Gymnophiona of India: A Taxonomic Study.
     Rec. Zoological  Survey  India, Occasional Paper No. 172: 1-126.

Pough, F.H., R.M. Andrews, J.E. Cadle, M.L. Crump, A.H. Savitzky, K.D. Wells. 1998.
     Herpetology. Prentice Hall Press, New Jersey.

Taylor, Edward H. 1968. The Caecilians of the World: A Taxonomic Review. University    
     of Kansas Press, Lawrence.


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