Z. 2001. Syst. Evo1ut.-forsch. 29 (1991)304-311 Received on 18.

January 1991
0 1991 Verlag Paul Parey, Hamburg und Berlin
ISSN 0044-3808

Department of Geology, University of Brictol, England

Adult tooth crown morphology in the Typhlonectidae

(Amphibia:Gymnophiona)
A reinterpretation of variation and its significance
By M. WILKINSON

Abstract
Adult tooth crown morphology is surve ed in the caecilian family Typhlonectidae. All adult
typhlonectids have recurved monocuspedl teeth that ma bare blade like lateral flanges. Teeth
in sizes, curvature and the development of lateral ianges within individuals, both within
lirbetween dental series. Most species have a pointed tooth crown, but the teeth of Typhlonectes
compressicaudus have broadly dilated crowns, which show some ontogenetic variation in their
elaboration. The teeth of Potomotyphlus kaupii are distinctly narrow, relatively more numerous
and more closely placed within series than in other typhlonectjds., The functional, taxonomic
and ph logenetic significance of the observed interspeci ic variation is discussed. Considerations
of the gehaviour of captive specimens and of morphology indicate that dilated tooth crowns are
probably not ada tations for scraping as had been suggested. The teeth of I! kaupii may be
adapted for handl?ng small pre Previous contradictory reports of typhlonectid tooth crown
morphology are interpreted as t l e result of misidentification of specimens.
Key words: Gymnophiona - Typhlonectidae - Dentition - Feeding - Taxonomy

Introduction
Caecilians typically have pointed tooth crowns with either one or two cusps (WAKEand
WURST1979). An interesting exception to this, within the aquatic genus Typblonectes,was
mentioned by TAYLOR (1968) and more fully described by WAKE(1978) and WAKEand
WURST(1979). TAYLOR (1968) noted that the teeth of the holotype of I: obesus Taylor were
somewhat spatulate rather than pointed. WAKE(1978) confirmed this unusual morphology
in a further series of specimens assigned to I: obesus and compared the teeth of this form
with the more typical pointed teeth of a congeneric species identified as I:compressicaudus
Dumeril and Bibron. She argued that the observed morphological differences might be
correlated with differences in diet between these species and that, whereas caecilians are
generally believed to be generalist predators, some species might be adapted for dietary
specialisation. WAKEand WURST(1979) provided scanning electron micrographs (SEMs)
of representative teeth of I: compressicaudus and I: obesus and further corroborated the
differences reported by WAKE(1978). GREVEN (1986) published SEMs of the teeth of
Tcompressicaudusshowing distinctly spatulate tooth crowns, but he offered n o discussion
or explanation of the discrepancy between his report of tooth crown morphology of this
species and the reports of WAKE(1978) and WAKEand WURST(1979).
I n this paper I briefly survey variation in adult tooth crown morphology within the
Typhlonectidae and attempt to resolve the conflicting reports of the tooth crown morpho-
logy of Typhlonectes compressicaudus. The taxonomic and functional significance of this
variation is discussed.

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 Adult tooth mown morphology in the Typhlonectidue 305

Materials and methods

Teeth were observed in situ, with a binocular dissecting microscope, in alcohol preserved speci-
mens. Representatives of all species o f t phlonectid recognised by NUSSBALJM and WILKINSON
(1989), with the exception of Nectocaecilu huydee (Roze), and type material representing these
s ecies and associated synonyms were examined. Selected single tooth crowns were extracted,
creaned manually, gold plated and observed with a scannin electron microsco e. Figures of teeth
were prepared from SEMs, and those of typhlonectid heats from camera luci$a sketches. Obser-
vations were made on the feeding of captive Tjphlonectes compressicaudus and T. nutuns (Fischer)
and on the terrestrial locomotion of T. nutans.

Results
All typhlonectids have monocuspid recurved teeth. There is minor variation in size
(TAYLOR 1968) and the degree of curvature both within and, more obviously, between
tooth series (Fig. 1). In all series there is an antero-posterior decrease in element size. The
curvature of teeth in more posterior loci tends to be skewed, such that these teeth project
more posteriorly than they would if their curvature were restricted t o a plane perpendicu-
lar t o the tooth series. Rather than having a simple conical form, typhlonectid teeth may
have blade-like lateral flanges of variable extent. In this respect, typhlonectid teeth resem-
ble the dentary teeth of the caeciliaid genera Dermophis and Gymnopis described by WAKE

Fig. 1. Lingual views of typhlonectid

teeth redrawn from SEMs. - A: an- t d
terior premaxillary of Chthonerpeton
indistincturn (Bar = 200 p m ) ;B: right
anterior dentary of Potomotyphlus
kaupii (Bar = 150 pm). Note the asym-
metric development of lateral flanges,
gentle skewness and relative narrow-
ness; C: anterior dentary of Typhlonec-
tes nutuns (Bar = 231 pm); D: an-
terior vomeropalatine of Z nutuns (Bar
= 136 pm). Note the poor development
of of lateral flanges; - E-H: ?: compres-
sicuudus showing ontogentic, post,
itonal, and serial variation; E: an-
terior dentary of newborm (Bar =
136 pm); F: anterior dentary of adult
(Bar = 200 p m ) ; G: more posterior
dentary of adult showing skewness
(Bar = 176 p m ) ; H: anterior right
premaxillary (Bar = 176 pm)
306 M.Wilkinron

and WURST(1979). In these genera and in typhlonectids the flanges arise some distance
distal to the tooth base, but in Demophis and Gymnopis the lateral flanges of the dentary
teeth appear to be expanded proximally to a point and then have relatively straight edges
converging to a point distally more rapidly than the more subconical ‘body’ of the teeth
which gives the teeth a distinctive arrowhead o r diamond shape. Lateral flanges are more
uniformly developed along the proximal-distal axis of the teeth of typhlonectids and con-
verge distally at approximately the same rate as the ‘body’ of the teeth. In typhlonectids
anterior teeth have their flanges developed symetrically whereas at more posterior loci the
flanges are asymmetric. The asymmetry develops gradually with the reduction in relative
size of the anterior lateral flange of each tooth and continues posteriad such that the an-
terior flange may be more or less absent in the posteriormost teeth. Of the four series of
teeth common to all typhlonectids, lateral flanges are best developed on the relatively large
teeth of the dentary series, are less extensive on the slightly smaller teeth of the premaxil-
lary-maxillary series and are least apparent on the smaller teeth of the splenial and vomero-
palatine series.
Pointed teeth conforming to the above description typify all species of Chthonerpeton
Peters, Nectocaecilia Taylor, and Potomotyphlus Taylor. Among these forms, the teeth of
Potomotyphlus are highly distinctive because of their exceptional narrowness and very well
developed lateral flanges (Fig. 1b). The teeth of Potomotyphlus are also more numerous
(TAYLOR 1968) and are positioned more closely together into series than in other
typhlonectids.
Within Typhlonectes Peters I find, with TAYLOR (1968), WAKE(1978) and WAKEand
WURST(1979), that there are two distinct tooth crown forms. Of the three species recog-
nised by NUSSBAUM and WILKINSON (1989) and WILKINSON (1989) two, Z eiselti Taylor
and 1: natans have the simple pointed tooth crowns that are characteristic of other
typhlonectids. The pointed tooth crowns of T. natans were first noted by FUHRMANN
(1914). In agreement with GREVEN (1986), and contrary to the reports of WAKE(1978) and
WAKEand WURST(1979), I find that T. compressicaudus, the type species of the genus, has
dilated, broadly spatulate tooth crowns. In addition, the type specimens of three species
described by TAYLOR (1968) from limited material (I:obesus, T. anguilkzformis and Nec-
tocuecilia ladigesi also have spatulate rather than pointed tooth crowns. The validity of
these latter three species names is in some doubt (see below).
There is considerable variation in the extent of dilation of the tooth crown and in the
development of lateral flanges in Typhlonectes compressicaudus. Much of this variation
appears to be ontogenetic. The most broadly dilated spatulate teeth are found in the short-
est and presumably youngest specimens. They are especially pronounced in newborns
(Fig. le). With increasing size and age, and the replacement of teeth, the dilation of the
crown is reduced but remains distinctive (Fig. If). Teeth of young animals also lack any
indication of the development of lateral flanges other than the dilated tooth’crown itself
which has blade like lateral margins. In teeth from older specimens there may be indica-
tions of weak lateral flanges along the body of the teeth that are continuous with the
lateral edges of the dilated tooth crowns. Some differences exist both between teeth of
different series and within the same series in this species. Size differences between series
and the skewness of posterior elements within series parallel these features in other
typhlonectids. In addition, teeth of the dentary series have the most dilated crowns. The
smaller teeth of the other series haver less broad crowns and may be rather chisel shaped
(Fig. Ih).
 Adult tooth crown morphology in the Typhlonectidae 307

Discussion
Taxonomic considerations
In order to resolve the contradictory reports of typhlonectid tooth crown morphology it
is first necessary to consider some problems in the taxonomy of the Typhlonectidae. In
the last monographic taxonomic treatment of the Typhlonectidae, TAYLOR (1968) included
six species in the genus Typhlonectes. Of these only three, T. rompressicaudus, I: eiselti and
T. natans, can be considered to be of certain validity (WILKINSON 1989). The two nominate
species T. anguilkzformis and T. obesus, the type specimens of which have dilated tooth
crowns, were established by TAYLOR largely on the basis of characteristics of body shape,
the former being relatively slender and the latter being relatively stout bodied. In T. com-
pressicaudus and 7: natans, which are well represented in scientific collections, there is con-
siderable variation in the proportions of the body with some specimens approaching the
slender form that TAYLOR (1968) apparently considered diagnostic of 7: anguillafomis.
Indeed some specimens that TAYLOR (1968) assigned to 7: compressicaudus are
indistinguishable in terms of body shape and proportions from others that he later as-
signed to 1: anguillaformis (TAYLOR 1970) and to ‘I: obesus (TAYLOR 1973). None of the
specimens later assigned to I: obesus by TAYLOR (1973) are as extremely stout as is the
holotype of this nominate species. This difference is accounted for by the fact that the
holotype is a heavily gravid female and that its bodily proportions have no doubt been
affected by this condition (TAYLOR 1973). The ‘diagnostic’ stout body shape of 1:obesus is
therefore a characteristic that is restricted, among specimens assigned to this species, to
the holotype and is clearly due to ephemeral variation in reproductive state. The type of
N . ladigesi has a poorly expressed dorsal fin, which presumably led TAYLOR (1969) to de-
scribe it as a species of Nectocaecilia. The expression of the dorsal fin is now known to vary
greatly within Typhlonectes (WILKINSON 1988) and the fin of the type of N. kzdigesi falls
well within this range of variation. A series of features including, annular and vertebral
numbers, lung lengths, and the pattern of denticulations about the cloaca1 disc reliably
distinguish the valid forms T. compressicaudus, I: eiselti and 7: natans from one another. In
contrast the three nominate forms T. angillafomis, T. obesus and N.kzgigesi are indistin-
guishable from each other and from T. compressicaudus in regard to these features.
The present study grew out of the uncertainty surrounding the validity of Typhlonectes
anguillaformis, 1: obesus and Nectocaecilia ladigesi. TAYLOR’S (1968) report of a novel tooth
crown morphology in one of these nominate species and the subsequent corroborations
by WAKE(1978) and WAKEand WURST(1979) appeared to indicate a potentially useful
taxonomic character. My findings, which support those of GREVEN(1986), show that
dilated tooth crowns are a characteristic of I: compressicaudus that is also shared with speci-
mens assigned to 7: anguillaformis, I: obesus, and N . ladigesi TAYLOR (1968, 1970, 1973).
Variation in tooth crown morphology is therefore insufficient to distinguish I: compres-
sicaudus from these doubtful nominate species.
The geographic range of Typhlonectes compressicaudus encompasses the whole of the
Amazon Basin, from the eastern coast of Brazil to the Atlantic slopes of the Peruvian
Andes, and the rivers of the Guianan Shield. This extensive range subsumes the localities
from which the three doubtful nominate species have been reported. Further studies are
needed to discover whether any of the presumably allopatric populations that have been
variably assigned to 7: compressicaudus, 7: anguillaformis, 7: obesus, and Nectocaecilia
ladigesi, and that occur in the numerous tributaries and branches of the Amazonian and
Guianan drainage systems are sufficiently divergent as to warrant the recognition of dis-
tinct species.
Biochemical approaches to this taxonomic problem would be especially welcome. In
the absence of characters which serve to distinguish the above nominate species, the three
names 7: anguillaforrnis, I: obesus, and N . ladigesi should be regarded as junior synonyms
308 M . Wilkinson

of T. compressicaudus, and they have thus been excluded from the latest synopsis of the
Typhlonectidae (NIJSSBAUM and WILKINSON 1989).

Identification of specimens
The reports of pointed tooth crowns in Typhlonectes compessicaudus by WAKE(1978) and
WAKEand WURST(1979) are inconsistent with my observations and with the report of
GREVEN (1986). The teeth figured by WAKE(1978) and WAKEand WURST(1979) for this
species are essentially identical with those of T. natans. The anomalous reports may be
explained as a result of the misidentification of specimens. Specimens of 7: nutuns that
have been reported upon by WAKEin a series of papers prior to 1986, and which I have been
able to examine had been misidentified as T. compressicaudus and reported as such. Such a
misidentification is not particularly surprising because of the inadequate nature of avail-
able keys to differentiate the species of Typhlonectes. Z nutans can be distinguished from
T. compressicaudus by the presence of a geometrical pattern of nine, rather than ten, denti-
culations on the cloaca1 disc.
Phylogenetic significance
Given the predominance of pointed tooth crowns throughout the Gymnophiona, there
can be little doubt that the spatulate tooth crown form is a derived character state. As
such, it may be phylogenetically uninformative because of its restriction to a single
species. However, if further study were to reveal that 7: compressicaudus, as conceived
here, consists of a number of cryptic species, then the sharing of spatulate tooth crowns
would be evidence that such forms constitute a holophyletic assemblage to the exclusion
of other species to Typhlonectes.
Similarly, the distinctive narrow form of the teeth of Potomotyphlus is probably a
derived condition that is unique to this genus. TAYLOR (1968) included two species in
Potomotyphlus but it is presently considered monotypic (NUSSBAUM and WILKINSON 1989)
so that narrow tooth form is presently phylogenetically uninformative with regard to
species relationships.
The form of the lateral flanges of typhlonectid teeth might be characteristic of the
group but a more comprehensive knowledge of the distribution and form of the flanges in
non-typhlonectid caecilians is required before the attempt to extract phylogenetic infor-
mation from this character is warranted.
Tooth crown morphology therefore appears to provide no evidence that bears on the
hypothesised holophyly of the Typhlonectidae (NUSSBAIJM and WILKINSON 1989) or
hypothesised relationships within the family (WILKINSON 1989). Monocuspidness, how-
ever, is probably a derived condition within the Gymnophiona because bicuspedness is
widespread in other Lissamphibia. The distribution of monocuspedness might thus
delimit a holophyletic group of which the Typhlonectidae is a part. Monocuspid teeth are
found in some but not all genera of the Caeciliaidae (WAKEand WURST1979) and this
distribution SUPPOKS the hypotheses that the Caeciliaidae is not a ‘natural’ group (WAKE
and WURST1979) and is paraphyletic with respect to the Typhlonectidae (NUSSBAUM and
WILKINSON 1989). A parallel argument can be made concerning the presence of lateral
flanges which are also found in some but not all caeciliaids.

Functional significance
WAKE(1978) reported that the stomach contents of a series of juvenile ‘Typhlonectesobesus’
included fragments of wood and rock and the remains of insect, probably coleopteran,
pupae that, based on “the structure of the exoskeleton and other characteristics”, were
probably maintained in air rather than water. She argued that “7:obesus probably forages
at the water’s surface at least occasionally, in order to scrape prey items such as pupae from
 Adult tooth crown morphology in the Typhlonectidae 309

rocks and logs”. She further suggested that the aquatic typhlonectids “swim in a sine-wave
pattern not dissimilar from the terrestrial locomotor pattern, so it is possible that they
could move about o n rocks and vegetation at the sides and surface of the watercourse for
some time. Their dependence on water is secondarily derived and restrictions to that
habitat are not particularly evident”. Finally she concluded that the spatulate dentition is
related to scraping, suggesting a possible restriction to sedentary rather than freely moving
prey items. As additional support for this conclusion she pointed out that: 1. “teeth with
dilated crowns are well adapted for scraping”, 2. “the orientation of upper and lower teeth
so that they do not occlude could facilitate scraping from surfaces at different angles” and
3. similar dilated crowns characterise the deciduous foetal dentition of viviparous caeci-
lians which is believed to be employed in scraping epithelial cell secretions from the
oviduct surface (WAKE1977).
Several observations lead me to question WAKE’S (1978) conclusions. I have observed
the feeding of both captive Typhlonectes compressicaudus (with dilated tooth crowns) and
Tnatans (with the more typical pointed tooth crowns). Both these species show relatively
little inclination to feed at the water’s surface in captivity, and feed more readily and su-
cessfully on the underwater surface of the substrate, often grasping prey items and pulling
them into burrows or other aquatic shelters. This behaviour may be homologous to the
similar feeding behaviour described by WAKE(1978) for terrestrial caecilians. O n land
these aquatic species are relatively clumsy and I am unaware of any report of their having
been captured or observed in the terrestrial environment in the wild. Typhlonectes tend to
have a richer supply of mucus associated with the skin than to terrestrial caecilians, and
out of water their mobility rapidly decreases as the skin dehydrates and the mucus appears
to gain adhesive properties which ‘glue’ the animal to the substrate. It is possible that these
species might be able to manouver themselves through relatively wet terrestrial habitat,
but prolonged emergence from the water, for feeding or for any other reasons, does not
seem particularly likely
Both Typhlonectescompressicaudus and I:natans can be maintained in captivity on a diet
of earthworms or on pieces of fish indicating that any dietary specialisation associated
with dilated tooth crowns would at most be facultative rather than obligate. MOODIE
(1978) reported that the diet of an Amazonian population of T. compressicaudus included
fish and shrimp indicating that this spatulate toothed form is fully capable of taking freely
moving rather than sedentary prey and does not eschew aquatic prey.
In addition, the recurved form and posterior orientation and position of the teeth of
adult caecilians suggests that, whatever their crown shape, they would nor be well suited
to scraping food items from substrates. This is particularly so in the case of typhlonectids
because the Typhlonectidae is characterised by a well developed rostral projection of the
snout beyond the mouth which, in combination with the orientation of the teeth, would
effectively preclude scraping (Fig. 2b). In foetal caecilians which probably do use their
teeth in scraping or rasping, the teeth are arranged in multiple rows on the external surface
of the mandible rather than within the buccal cavity This special feature together with the
relatively less prominent rostral projection of the snout of foetal typhlonectids (Fig. 2a)
allows for the possibility of scraping which would otherwise be impossible if the teeth
were entirely in the buccal cavity as they are in adults.
It seems unlikely that the possession of dilated tooth crowns in adult T compressicaudus
is associated with dietary specialisation as envisaged by WAKE(1978). The evolution of
adult dilated tooth crowns does not at present seem explicable in terms of adaptations of
the adult feeding system and could concievably be a relatively selectively neutral pleio-
tropic consequence of some other change. The dilated tooth crowns of Typhlonectes com-
pressicaudus adults are similar to those of foetal caecilians both of this and of related
(pointed toothed) species such as Chthonerpeton viviparum, Nectocaeciliz petersi (PARKER
and DUNN1961) and 7: natans (pers. observ.) although the tooth crowns of foetuses may
310 M.Wilkinson

. Fig. 2. Semidiagramatic illustration of head

sha e and tooth position in (A) foetal,and (B)
aduft typhlonectids. Note the array of foetal
teeth on surface of the mandible external to the
buccal cavity and the less prominent snout of
the foetus which may facilitate intraoviducal
feeding by ‘scraping’. The prominent snout of
the adult, the position of the adult dentition
entirely within the buccal cavity and the ost
erior recurvature of the adult teeth are alyfei
tures which indicate that ‘scraping’ is not a
mode of feeding that is likely to be employed
by adult typhlonectids (not to scale)
also be equiped with spikes. This suggests that dilated adult tooth crowns could be the
.result of a paedomorphic process which may also be indicated by the adult dentitional
ontogeny where replacement teeth have less broadly dilated crowns.
I n contrast, the narrow tooth form of Potomotyphlus may well be related to dietary
specialisation. Potomotyphlus is characterised by reduced choanal valve apertures, a dispro-
portionately small head and neck region, and by having more numerous teeth than in
other typhlonectids (TAYLOR 1968). The choanal valve flaps are partially fused and the
small aperture is concealed under a fleshy flap. The narrow teeth are more tightly packed
into series than in other typhlonectids. These features, together with the narrow tooth
shape, are undoubtedly derived within the Typhlonectidae.
Although there is no available information o n the feeding habits of Potomotyphlus, it
seems likely that feeding in this genus is specialised at least in terms of prey size. The
advantageousness of such specialisation might have been a cause of the evolution of
reduced head and gape size, but even if head size reduction evolved for some other reason,
change in the size range of prey that could be taken would have been an inescapable conse-
quence. It seems reasonable therefore to suppose that narrow teeth, tightly packed to-
gether into series evolved as adaptations facilitating the handling of small prey items. Re-
duction of the choanal valve aperture and its concealment are also interpreble as adapta-
tions related to feeding o n prey of small size. A reduced, concealed aperture would more
effectively prevent prey items from entering the nasal passages from the mouth. The
danger of prey escaping down the ‘wrong hole’ would be enhanced by the reduction in the
size of the prey
Acknowledgements
Most of the research reported here was carried out at the Museum of Zoolog University of
Michigan. I thank R. A. NUSSBAUM for his guidance and encouragement and aCo for access to
specimens in his care. I am grateful to M. H. WAKEfor allowing me to visit her lab and examine
s ecimens in her care. M. J. BENTON read and commented o n the manuscript during preparation.
$he following curators and institutions have also assisted my research by making specimens avail-
able on loan: C. W. MYERS,American Museum of Natural History; E. V. MALNATE, Academy of
Natural Sciences, Philadelphia; E. N. ARNOLD and B. T. CLARKE, British Museum of Natural
History; R. C. DREWES,California Academy of Sciences; C. J. McCoy, Carnegie Museum;
H. MARX,Field Museum Natural History; W. E. DUELLMAN, University of Kansas, Museum of
Natural History; J. WRIGHT,Los Angeles County Museum; J. ROSADO,Museum of Compara-
tive Zoology, Harvard; V. MAHNERT and J.-L. PERRET, Museum d’Histoire Nature1 de Geneve;
J. LESCURE,Museum National d’Histoire Naturelle, Paris; 0. ELTER,Museo di Zoologia,
Torino; F. TIEDEMANN, Naturhistorisches Museum, Wien; W. BOHME, Zoologisches For-
schungsinstitut und Museum Alexander Koenig, Bonn; R. GUNTHER,Humboldt-Universitat,
Zoologisches Museum, Berlin; H. W. KOEPKE and H. WILKENS, Universitat Hamburg, Zoolo-
isches Museum; J. RASMUSSEN, Zoologisk Musem, Copenhagen; U. GRUBER,Zoologische
!taatssammlung, Miinchen.
Thanks are due to D. SENNand R. KURTHof the Zoologisches Institut, Universitat Basel and
to M. DELSOLand J.-M. EXBRAYAT of the Faculti Catholique des Science de Lyon for providing
the opportunities to observe living Typhlonecter compessicaudus. This work was supported in
part by SERC grant GR/F 87912.
 Adult tooth crown morphology in the Typhlonectidae 311

Summary
The majority of typhlonectids have pointed tooth crowns as is typical of caecilians. Typhlonectid
teeth are monocuspid. Serial variation in size, skewness and the development of lateral flanges,
and differences in tooth sizes in different series characteries individuals of all typhlonectid
species. 5 hlonectes compressicaudus is exceptional in having broadly dilated tooth crowns. Con-
fusion in t f e literature over the shape of the teeth of T. compressicaudus has been due to the mis-
identification of specimens. T. obesus, T. anguillaformis and Nectocaecilia ladigesi, the type speci-
mens of which have dilated tooth crowns, are best regarded as junior synonyms of I: com res
sicaudus. The functional si nificance of dilated tooth crowns is discussed. It is unlikely that ties;
teeth are associated with feeding or dietary specialisations and no adaptive si nificance can be
ascribed to dilated tooth crowns in adult caecilians at present. Potomoty hfus has relatively
P
numerous, distinct1 narrow teeth that are interpreted as components o as feeding system
adapted for the handing of small sized prey.

Zusamrnenfassung
Die Morphologie der Zahnkronen adulter Typhlonectidae (Amphibia: Gymnophiona).
Eine Reinterpretation der Variation und ihrer Bedeutung
Erwachsene Typhlonectidae haben einwarts gebogene monocus ide Zahne, an denen flache Sei-
tenflachen vorhanden sein konnen. Groge, Kriimmung und Ausiildung der Seitenflachen variie-
ren beim gleichen Individuum sowohl innerhalb als auch zwischen den Zahnreihen. Die meisten
Arten haben spitze Zahnkronen, aber die Zahne von Typhloneaes compressicaudus besitzen ver-
breiterte Kronen, deren Ausgestaltung wahrend der Ontogenese variiert. Die Zahne von Poto-
motyphlus kaupii sind deutlich schmaler, relativ zahlreicher und enger beieinander stehend als bei
anderen Typlonectiden. Die funktionelle, taxonomische und phylogenetische Bedeutung der Ei-
genarten wird erortert. Typhlonectes obesus, 7: anguillaformis und Nectocaecilia ladigesi werden
1:com ressicaudus als Synonyme zugeordnet. Verhaltensbeobachtungen fuhren im Verein mit der
Mor fologie zu der Auffassung, da8 verbreiterte Zahnkronen nicht als eine Anpassung an Ab-
schagen von Nahrung zu werten ist, was angenommen wurde. Die Zahne von I! kaupiz konnen
als Anpassung zur Bewaltigung kleiner Beute elten. Fruhere widerspruchliche Betrachtungen
iiber die Zahnkronen der Typhlonectidae werfen durch falsche systematische Zoordnung von
Individuen interpretiert.
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Author’s address: MARKWILKINSON,Department of Geology, University of Bristol, Queens
Road, Bristol BS8 1RJ, England