Gymnophiona

▪ amphibian

Introduction

also called  Apoda 

 one of the three major extant orders of the class Amphibia. Its members are known as caecilians (caecilian), a name derived from the Latin word caecus, meaning “sightless” or “blind.” The majority of this group of wormlike amphibians (amphibian) live underground in humid tropical regions throughout the world. Because of their relatively hidden existence, caecilians are unfamiliar to the layperson and are not usually considered in discussions about amphibians. They are nevertheless a fascinating group of highly specialized amphibians about which there is still much to be learned.

General features

Size and range

      Several species of caecilians (caecilian) in the South American genus Caecilia exceed 1 metre (about 3.3 feet) in total length; the largest known caecilian is C. thompsoni, at 152 cm (about 60 inches). The smallest caecilians are Idiocranium russeli in West Africa and Grandisonia brevis in the Seychelles; these species attain lengths of only 98–104 mm (3.9–4.1 inches) and 112 mm (4.4 inches), respectively.

Distribution and abundance

      Caecilians are found in tropical areas throughout the world. Of the three families that inhabit southern Asia, one occurs in geographically separated pockets in India, Southeast Asia, Borneo, and the Philippines. Three families are widespread in the American tropics, but only two are endemic; two families occur in Africa, and one is endemic. On the Seychelles there are three genera native to the islands, although caecilians are not found on any other islands in the Indian Ocean. None have been found on Madagascar or New Guinea. Approximately 170 caecilian species are known to exist, and up to 5 species have been found to inhabit the same area in the Amazon Rainforest.

Natural history

Breeding behaviour

      Information about annual reproductive (reproduction) patterns among caecilians (caecilian) is limited. The breeding period of some Asiatic ichthyophiids seems to be aseasonal or at least without seasonal constraints. At least one species, Ichthyophis glutinossus in Sri Lanka, mates only during the rainy season. Females of viviparous (viviparity) species have a biennial reproductive cycle; the viviparous Dermophis mexicanus in Guatemala mates in the early part of the rainy season, and gestation takes one year.

      All caecilians are believed to have internal fertilization. This is achieved by means of the phallodeum, a copulatory organ in males that is modified from the cloacal wall (cloaca). Eggs (egg) of all members of the families Ichthyophiidae and Rhinatrematidae are deposited in burrows in mud that is close to water. The females watch over these clutches, which may hold up to 54 eggs. Upon hatching, the larvae (larva) leave the burrows to make their homes in ponds and streams. Some caecilians deposit eggs on land, and in different species these hatch as larvae or small adults. Three families have viviparous species to which usually no more than four young are born at one time. Aquatic typhlonectids are viviparous and produce larvae. The caecilian fetus emerges from the egg membrane as soon as its meagre yolk supply is exhausted; it uses its deciduous teeth, adapted for scraping, to obtain secretions and epithelial tissues (epithelium) from the oviduct lining.

Feeding habits

      The diet of terrestrial caecilians is mainly earthworms (earthworm) and other soft-bodied prey. Feeding either aboveground or in subterranean burrows, terrestrial caecilians are believed to locate their quarry by means of a chemosensory tentacle on each side of the head. They capture their prey (predation) with their powerful recurved teeth, masticate, and swallow. Aquatic caecilians, the typhlonectids, prey on fishes (fish), eels (eel), and aquatic invertebrates (invertebrate).

Form and function

 Caecilians have long, limbless, cylindrical bodies that abruptly end behind the cloaca or short tail. Annuli (primary grooves) in the skin encircle the body and form segments; in some taxonomic groups, secondary and tertiary grooves partially circumscribe the body. Within the tissue of the annuli, bony scales (scale) of dermal origin usually occur. The heads of caecilians are blunt, and their skulls are bony and compact. Centres of ossification have fused, which has reduced the number of independent cranial bones in caecilians in comparison with anurans (Anura) and salamanders (salamander); for example, a single bone, the os basale, forms both the floor of the braincase and the posterior part of the skull. Teeth are found on all jaw bones, and a palatal series of teeth appears in addition, medial to the maxillary series. A U-shaped facet, which articulates with the quadrate and also has a long retroarticular process that serves as an attachment site for three major jaw muscles, is located on the lower jaw. The vertebral column is made up of an atlas (the first vertebra of the neck) and 95 to 285 trunk vertebrae; no differentiated sacral vertebrae are present. Double-headed ribs are found on all vertebrae except the atlas and the terminal three to six vertebrae. Of the three amphibian orders, only caecilians have an axial musculature in which all the hypaxial components, excluding the subvertebral musculature, form an outer muscular sheath. This sheath, which is anchored to the skin by fibrous connective tissue, is all but disconnected from the vertebral musculature and thereby allows the skin and superficial muscles to move as a single unit. The degenerate eyes are covered with bone or skin. These adaptations make it possible for the caecilian to feed, reproduce, and avoid enemies within their subterranean realm. The features of aquatic caecilians of the family Typhlonectidae are representative of secondary adaptations.

Classification

Distinguishing taxonomic features

      The families of caecilians (caecilian) are distinguished from one another by morphological characters and features of their life histories. The Jurassic (Jurassic Period) fossil caecilian has small limbs, whereas all living caecilians are limbless. The living families are distinguished from one another by the presence or absence of a tail, amount of fusion of cranial bones, degree of kineticism of the skull, nature of the annular grooves, and structure of the phallodeum. Aquatic larvae are present in three families; in the other families, larvae (larva) are absent.

Annotated classification

      The classification below is based on the work of Ronald A. Nussbaum and Mark Wilkinson (1989).

Order Gymnophiona (Apoda)

 Elongate limbless amphibians well adapted for burrowing; body segmented by annular grooves, with some species containing scales; tail, if present, short and pointed; skull compact, in which some elements are fused; eyes small, covered with skin or bone; teeth curved; left lung often primitive; projectable sensory tubercle between eye and nostril; protrusible copulatory organ in males; aquatic larvae (with gill slits but no external gills), direct development of terrestrial eggs, or viviparous; about 170 species.

      Family Caeciliidae

 Paleocene (65.5–55.8 million years ago) to present; tail absent; mouth recessed; premaxillae fused with nasals; prefrontals absent; squamosal articulating with frontal; usually no aquatic larval stage; 23 genera, 90 species; adult size 10–152 cm (4–60 inches); South and Central America, Africa, India, and Seychelles.

      Family Ichthyophiidae

 No fossil record; tail present; mouth subterminal (partially recessed); premaxillae not fused with nasals; prefrontals present; squamosal articulating with frontal; aquatic larvae; 2 genera, 37 species; adult size 40–50 cm (16–20 inches); Southeast Asia, peninsular India, Sri Lanka, Sumatra, Borneo, and Philippines.

      Family Rhinatrematidae

 No fossil record; tail present; mouth terminal; premaxillae not fused with nasals; prefrontals absent; squamosal not articulating with frontal; aquatic larvae; 2 genera, 9 species; adult size 25–32 cm (10–13 inches); South America.

      Family Scolecomorphidae

 No fossil record; tail absent; mouth recessed; premaxillae not fused with nasals; prefrontals present; squamosal not articulating with frontal; no aquatic larval stage; 2 genera, 7 species; adult size 40–45 cm (16–18 inches); Africa.

      Family Typhlonectidae

 No fossil record; tail absent; mouth recessed; premaxillae fused with nasals; prefrontals absent; squamosal articulating with frontal; no larval stage; adults aquatic; 4 genera, 18 species; adult size 50–72 cm (20–28 inches); South America.

      Family Uraeotyphlidae

 No fossil record; tail present; mouth recessed; premaxillae not fused with nasals; prefrontals present; squamosal articulating with frontal; aquatic larvae; 1 genus, 4 species; adult size 28–30 cm (11–12 inches); southern peninsular India.

Critical appraisal

      The present classification is strongly supported by morphological and biochemical evidence, except for family Caeciliidae, which contains groups of unrelated species.

Additional Reading

Edward H. Taylor, The Caecilians of the World: A Taxonomic Review (1968), is the first major review of caecilians, and it served to ignite the interest of other scientists. Similarly, William E. Duellman and Linda Trueb, Biology of Amphibians (1986, reissued 1994), offers the first general review of caecilians' biology and anatomy. A more recent and less detailed review of caecilians is available in George R. Zug, Laurie J. Vitt, and Janalee P. Caldwell, Herpetology: An Introductory Biology of Amphibians and Reptiles, 2nd ed. (2001). The most comprehensive review of caecilians is Werner Himstedt, Die Blindwühlen (1996). The current classification is derived from Ronald A. Nussbaum and Mark Wilkinson, “On the Classification and Phylogeny of Caecilians (Amphibia: Gymnophiona), a Critical Review,” Herpetological Monographs, 3:1–42 (1989).William E. Duellman

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