© 2010 Benjamin Tapley (1 of 1)
Diagnosis: A key to Indian caecilian genera is presented in Bhatta (1998); Indian species belonging to the genera Ichthyophis and Uraeotyphlus can be distinguished by having a pointed tail and longitudinal vent (vs. rounded tail and transverse vent in species of the genera Gegeneophis and Indotyphlus). Laterally striped forms with pointed tails and longitudinal vents belong to the genus Ichthyophis.
Ichthyophis beddomei is a small species of Ichthyophis with a cream or yellow lateral stripe on the body. It can be distinguished from other striped Ichthyophis of the Western Ghats by the following combination of characters: lacking a whitish midventral stripe, having fewer than 320 annuli, a tentacle almost as close to the naris as to the eye, a narrow and pointed head, a broad lateral stripe that is distinctly evident on the mandible, and an inner mandibular tooth row not much shorter than the dentary row (Wilkinson et al. 2007).
Description: This caecilian species reaches a total length of 170-275 mm (Pillai and Ravichandran 2005), with a short and broad body (Bhatta 1998). It typically has 260-295 primary and secondary annuli (Pillai and Ravichandran 2005). Anterior annuli have one to two rows of dermal scales, increasing to six to eight rows of scales toward the posterior (Pillai and Ravichandran 2005). The snout is relatively long and somewhat pointed, and projects slightly over the jaw (Bhatta 1998; Pillai and Ravichandran 2005). The nostrils are nearly terminal but are just barely visible when the head is viewed from above (Bhatta 1998). The tentacles are long and conical (Bhatta 1998), and the tentacular aperture is near the upper lip and about equidistant between the nostril and the eye (Bhatta 1998; Daniels 2005). The eye rests in a semicircular socket and is quite distinct (Taylor 1968). Typical dentition consists of 19-19 premaxillary-maxillary teeth, 22-23 prevomeropalatine, 20-20 dentary, and 22-22 splenial teeth (Pillai and Ravichandran 2005). The collars are wider than the head and are fused dorsally (Pillai and Ravichandran 2005). Behind the jaw, the first nuchal groove is distinct ventrally and dorsally, while the second nuchal groove is only distinct ventrally; the third groove, meanwhile, is hard to distinguish from the rest of the annuli and does not cross the midventral line (Pillai and Ravichandran 2005). The second collar groove has two well-defined folds on the dorsal side (Bhatta 1998). This species has a short pointed tail with five to nine folds (Daniels 2005). The vent is longitudinal with lobulate denticulations around it: three anterior to the vent, and four on each side (Pillai and Ravichandran 2005). The tip of the tail is compressed laterally and has no unsegmented terminal shield (Pillai and Ravichandran 2005).
Body color is violet brown dorsally and light brown ventrally, with a characteristic yellow or cream stripe running laterally from the snout to the tail tip (Bhatta 1998; Pillai and Ravichandran 2005). The yellow stripe often widens at the first collar, but the configuration of the stripe on the collars is variable (Pillai and Ravichandran 2005). The area around the vent is distinctly yellow (Daniels 2005).
Similar species: I. beddomei can be distinguished from similar striped species of Ichthyophis in the Western Ghats in the following ways. It differs from Ichthyophis longicephalus by having a tentacle almost as close to the naris as to the eye (vs. tentacle more than 2x as far from the naris than from the eye in I. longicephalus; Wilkinson et al. 2007), by having fewer than 320 annuli (vs. more than 320 annuli in I. longicephalus; Wilkinson et al. 2007), by lacking yellow patches at the sides of the second collar, first collar and jaw angles and lacking a longitudinal midventral line (vs. present in I. longicephalus; Bhatta 1998), and having the tentacle further from the eye (tentacle-naris distance/tentacle-eye distance ratio<1.25, vs. TN/TE>1.75 in I. longicephalus; Wilkinson et al. 2007); from I. tricolor by lacking a whitish midventral stripe (vs. present in I. tricolor; Wilkinson et al. 2007); from I. kodaguensis by having the tentacle relatively closer to the eye (TN/TE < 1.25, vs. TN/TE of 1.25-1.58 in I. kodaguensis; Wilkinson et al. 2007), by having a narrow pointed head (vs. broad and round in I. kodaguensis), by having a broad lateral stripe on the mandible (vs. indistinct narrow lateral stripe on the mandilble in I. kodaguensis), and by having an inner mandibular tooth row not much shorter than the dentary row (vs. inner mandibular tooth row much shorter than dentary row) (Wilkinson et al. 2007).
Distribution and Habitat
Life History, Abundance, Activity, and Special Behaviors
Behavior of wild-caught individuals was regularly observed in captivity over a period of two years by Bhatta (1999). Individuals were reared in a terrarium with a 15 cm thick substrate of moist humus-rich soil taken from the locality where the animals were captured (and changed periodically), and fed with locally obtained earthworms once a week. This species was most active at night, preferring to rest inside burrows during the day. At night animals emerged to feed on the soil surface, sometimes emerging only partially with just the head protruding out of the tunnel and retreating quickly if disturbed. New burrow openings appeared to be created only if existing burrow openings had been blocked, suggesting that this species prefers to use existing burrows rather than regularly creating new ones. Tunnels were found to be coated with mucus. When males and females were put into a terrarium without any soil substrate, several instances of both attempted and apparently completed copulation were observed. Males encountering another individual moving in the opposite direction positioned their ventral surface against that of the other individual, with contact only at the posterior of the animals; no twisting or whole-body contact was observed. If the partner was female, cooperation ensued and she allowed phallodeum insertion by the male, with two or three attempts sometimes required to achieve a successful copulation.
I. beddomei is an oviparous species (Gower et al. 2004), breeding between March and December (Daniel 2002). It lays its eggs near water in a clutch bound together by filaments (Daniels 2005). Clutches have been collected from stream banks during January to April (Bhatta 1999). Clutches consist of about 15 eggs (Daniels 2005), and range from 6-29 eggs (Bhatta 1999), with Tikader (1964) also providing a report of a clutch with 12 eggs. The eggs are spherical and are about 1.2 cm in diameter (Daniels 2005). The mother exhibits parental care by coiling around the eggs and protecting them during development (Daniels 2005). Bhatta (1999) also reported that captive I. beddomei females did not seem to distinguish their own clutch from those of other females, and abandoned the clutches at later stages of development (in agreement with the observations of Balakrishna et al. 1982, who reported finding late-stage unattended clutches that hatched soon after being brought into captivity). Mothers keep the clutches moist, rotate the eggs, and may move them around (Bhatta 1999). The aquatic larvae develop rapidly and lose their gills in about two days, after which time they leave the water and burrow into the soil (Bhatta 1999; Daniel 2002).
Oogenesis and folliculogenesis have been extensively studied in Ichthyophis beddomei. This species appears to have an annual cycle, reproducing once a year (Masood-Parveez and Nadkarni 1993a). Ovarian morphology and histology were studied in specimens collected from Sringeri, Karnatak State during the June-August monsoon, and maintained in captivity. The ovarian activity has three phases: (1) pre-breeding (August-November), where the ovarian mass increases, follicles undergo vitellogenesis, and the percentage of atretic follicles decreases; (2) breeding (December-February), characterized by a high number of large vitellogenic follicles, a low number of pre-vitellogenic follicles, and dividing oogonia; (3) post-breeding (March-July), characterized by a decreased follicle diameter, an increased percentage of atretic follicles, a decreased percentage of vitellogenic follicles, and the presence of post-ovulatory follicles (Masood-Parveez and Nadkarni 1993a).
Ovarian morphology is similar to that of other caecilians (Masood-Parveez and Nadkarni 1993a, b; Wake 1968). The germinal epithelium is distributed along the length of the ovary in a segmental arrangement, with alternation of the germinal zone and the follicular zone (Masood-Parveez and Nadkarni 1993a). The structure of the follicle at all stages (pre-vitellogenic, vitellogenic, and post-ovulatory) is similar to that of other amphibians (Masood-Parveez and Nadkharni 1993b). I. beddomei has two types of follicular atresia, one resembling those of other vertebrates (fish, pigeon, lizard, some mammals) where the oocyte shrinks and the follicular layer is left behind, and the other resembling those of reptiles and anurans (a different lizard species and one toad species), where granulosa hypertrophy and hyperplasia occurs (Masood-Parveez and Nadkarni 1993b).
Masood-Parveez and Nadkarni (1991) described the annual cycle of oviductal changes in this species. During the first part of the post-breeding season (March-April) oviducts resemble threads. From August to February, they are enlarged and contain three distinct morphological structures: the anterior third of the oviduct, the pars recta, consists of a straight thin tube; the middle third forms a convoluted tube, called the pars convoluta; the posterior third consists of a straight, thick tube, the pars uterina, which joins the cloaca. Glandular cells in the pars recta and the pars convoluta hypertrophy during October to February. In the pars convoluta and the pars uterina, epithelial cells develop cilia from September/October to February. In the pars uterina, glandular cells hypertrophy during August to February. Oviductal mass was maximal between October to February. Thus, oviductal activity is coordinated with ovarian activity. Unlike anurans and urodeles, caecilians, as exemplified by this species, have glands in all three regions of the oviduct (Masood-Parveez and Nadkari 1991).
Adrenal (interrenal) activity peaks during the breeding phase (January and February) and also in the post-breeding phase (monsoon season in June and July) (Masood-Parveez et al. 1992). The interrenal tissue in this species is divided morphologically into four zones. The first zone is similar to that of teleosts, with interrenal islets on the kidneys attached to the dorsal aorta, while islet structure in the second, third, and fourth zones resembles that of urodeles. As is the case for adrenocortical cells of anurans and urodeles, I. beddomei interrenal adrenocortical cells appear to have steroidogenic activity. It has been speculated that the peak in adrenocortical activity seen during the non-breeding season might function to maintain osmotic balance during heavy monsoon rains, when they are likely to be flooded out of their burrows (Masood-Parveez et al. 1992).
Masood-Parveez et al. (1994) note that pituitary glands also undergo morphological changes that are coordinated with the breeding cycle of I. beddomei. During the pre-breeding phase in August to November, pituitary B2 cells undergo an increase in nuclear diameter and degranulation of the cytoplasm. B2 cells are thought to be involved in secreting gonadotropins in anurans and urodeles as well. Pituitary B3 cells show an increase in nuclear diameter during November to January, correlating with increased activity of adrenocortical cells. The B3 cells secrete corticotropins in anurans and urodeles.
The neuroanatomy of I. beddomei with respect to gonadotropin-releasing hormone resembles that of many anuran and urodele species. There are two distinct areas in the brain expressing GnRH; one in the forebrain and terminal nerve, which contains GnRH with high sequence similarity to mammalian GnRH, and the other in the midbrain tegmentum (dorsal pallium), with high sequence similarity to chicken GnRH-II. The distribution of mGnRH-expressing neurons in I. beddomei is intermediate between the state for salamanders and frogs. In I. beddomei, mGnRH neurons are located within the terminal nerve, the midbasal telencephalon, and the septopreoptic area. In urodeles, mGnRH neurons are enriched in the intercranial segment of the terminal nerve, whereas in anurans, mGnRH neurons are enriched in the septopreoptic area, and only occasionally found in the terminal nerve (Pinelli et al. 1997).
The presence of cGnRH-II in the dorsal pallium has been described for Ichthyophis beddomei, a gymnophione, and Taricha granulosa, a urodele, but not for adult anurans. However, cGnRH-II is present in the dorsal pallium for the frog Rana esculenta during the last few days of metamorphosis, as well as in the dorsal forebrain of juvenile lizards (Podarcis sicula). Presence of cGnRH-II in the dorsal pallium may therefore be a primitive character (Pinelli et al. 1997).
Trends and Threats
Possible reasons for amphibian decline
General habitat alteration and loss
Ichthyophis beddomei has been found to be the sister taxon to I. tricolor (Gower et al. 2002).
The species was first described by Peters (1879), who named it after Colonel Richard Henry Beddome, a British officer and naturalist in the Indian Forestry Department (Taylor 1968). The name has also been spelled as Ichthyophis beddomii in older literature (e.g., Daniel 2002).
Footage shot by the scientists of WGRC/ZSI, Kozhikode, in Kerala, India.
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Written by Omar Malik and Kellie Whittaker (oam AT berkeley.edu), UC Berkeley
First submitted 2010-02-02
Edited by Kellie Whittaker; updated Ann T. Chang (2013-09-09)
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