Gymnophiona

Amphibian

WRITTEN BY
William E. Duellman
Curator, Division of Herpetology, Museum of Natural History; Professor of Systematics and Ecology,
University of Kansas, Lawrence. Author of The Hylid Frogs of Middle America.
www.britannica.com

Gymnophiona, also called Apoda, one of the three major extant orders of the class Amphibia. Its
members are known as caecilians, a name derived from the Latin word caecus, meaning “sightless”
or “blind.” The majority of this group of limbless, wormlike amphibians 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 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 10 known families, 5 occur in the
Americas, whereas Africa and mainland Asia harbour 3 families each. Caecilians are also found in
Indonesia, Sri Lanka, the Philippines, and the Seychelles. On the Seychelles there are three genera
native to the islands, although caecilians are not found on any other islands in the Indian Ocean. No
caecilians have been found on Madagascar or New Guinea. Approximately 180 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 patterns among caecilians 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 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. Eggs 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 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 from the oviduct lining.

Feeding habits
The diet of terrestrial caecilians is mainly earthworms 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 with their
powerful recurved teeth, masticate, and swallow. Aquatic caecilians, the typhlonectids, prey
on fishes, eels, and aquatic invertebrates.

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 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 and salamanders; 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 are distinguished from one another by morphological characteristics and
features of their life histories. The Jurassic 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 absent from most families.

Annotated classification

The classification below is based on the work of Ronald A. Nussbaum and Mark Wilkinson (1989),
Mark Wilkinson and others (2011), and Rachunliu G. Kamei and others (2012).

● ORDER GYMNOPHIONA (APODA)

NOTAR ACÀ QUE LOS CRITERIOS MORFOLÒGICOS MÀS IMPORANTES PARA

DIFERENCIAR ENTRE FAMILIAS SON, ENRE OTROS, SI TIENEN O NO COLA; PRESENCIA
O AUSENCIA DE CIERTOS HUESOS CRANEALES Y SI HAY FUSIÒN ENTRE ELLOS O NO
LA HAY; SI SON OVÌPAROS O VIVÌPAROS; SI HAY O NO ESCAMAS O ANILLOS DÈRMICOS
SECUNDARIOS, Y LA DENTADURA.

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 180 species.

o 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; 2 genera, 42 species; adult size 10–152 cm (4–60 inches); South and Central
America.

o Family Chikilidae
Jurassic (200–145.5 million years ago) to present; perforate stapes; septomaxillae and
prefrontals absent; lower jaws possess two rows of teeth; 1 genus, 7 species;
northeastern India.

o Family Dermophiidae
Cretaceous (145.5–65.5 million years ago) to present; secondary annuli and annular scales
present; viviparous; 4 genera, 13 species; Africa and Central and South America.

o Family Herpelidae
Cretaceous (145.5–65.5 million years ago) to present; perforate stapes (or stirrup bone) but
lack separate septomaxillae and prefrontal bone; 2 genera, 9 species; Africa.

o Family Ichthyophiidae
Cretaceous (145.5–65.5 million years ago) to present; tail present; mouth subterminal (partially
recessed); premaxillae not fused with nasals; prefrontals present; squamosal articulating with
frontal; aquatic larvae; 3 genera, 50 species; adult size 40–50 cm (16–20 inches); Southeast
Asia, peninsular India, Sri Lanka, Sumatra, Borneo, and Philippines.

o Family Indotyphlidae
Cretaceous (145.5–65.5 million years ago) to present; imperforate stapes and inner
mandibular teeth present with some teeth bicusped; viviparous forms lack scales and
secondary annuli; some forms are oviparous; 7 genera, 21 species; Africa, Seychelles, and
India.

o Family Rhinatrematidae
Cretaceous (145.5–65.5 million years ago) to present; tail present; mouth terminal; premaxillae
not fused with nasals; prefrontals absent; squamosal not articulating with frontal; aquatic
larvae; 2 genera, 11 species; adult size 25–32 cm (10–13 inches); South America.

o Family Scolecomorphidae
Jurassic (200–145.5 million years ago) to present; tail absent; mouth recessed; premaxillae not
fused with nasals; prefrontals present; squamosal not articulating with frontal; no aquatic larval
stage; adult stage without stapes and fenestrae ovales in the ear; 2 genera, 6 species; adult
size 40–45 cm (16–18 inches); Africa.
o Family Siphonopidae
Cretaceous (145.5–65.5 million years ago) to present; imperforate stapes and no inner
mandibular teeth; oviparous; 7 genera, 19 species; South America.

o Family Typhlonectidae
Cretaceous (145.5–65.5 million years ago) to present; tail absent; mouth recessed;
premaxillae fused with nasals; prefrontals absent; squamosal articulating with frontal; young
possess gills; no larval stage; adults aquatic; 5 genera, 13 species; adult size 50–72 cm (20–
28 inches); South America