Cecílias e Cobras Cegas

Reprint from:
Reproductive Biology and Phylogeny of Gymnophiona

(Caecilians)
Volume edited by Jean-Marie Exbrayat
Volume 5 of Series:
Reproductive Biology and Phylogeny
Series edited by Barrie G. M. Jamieson
Science Publishers, Enfield, NH, USA. 2006.
The layout of this reprint is not exactly as published (some lines at the end of some pages appear
at the beginning of the following page in the printed version) but the content is the same.
CHAPTER
2
Caecilian Phylogeny and
Classification
Mark Wilkinson1* and Ronald A. Nussbaum2
2.1 INTRODUCTION
Fifteen years ago, we published a critical review of caecilian phylogeny and
classification (Nussbaum and Wilkinson 1989). We hoped to establish some
stability in caecilian classification in the face of some highly divergent
phylogenetic hypotheses and alternative taxonomic treatments (Wake and
Campbell 1983; Duellman and Trueb 1986; Lescure et al. 1986; Laurent
1986). We concluded that caecilian phylogeny was too poorly known to
provide the basis for a working phylogenetic classification that recognized
only well-founded monophyletic groups. Instead, we provided an interim,
conservative classification in which 154 nominate species were partitioned
into six families with no sub- or suprafamilial ranks other than 34 genera.
Up until 1968, all caecilians were placed into a single family, the Caeciliidae,
and we recognized that the subsequent removal of distinctive subsets of
species in the establishment of additional families had left the Caeciliidae
a most likely paraphyletic assemblage of caecilians that did not fit into one
of the better circumscribed families. This is reflected in the relative numbers
of taxa: well over half of all recognized caecilian species and genera are
caeciliids.
Some important milestones have appeared over the last 15 years in
caecilian phylogenetics. Since our previous review, the first phylogenetic
study of the interrelationships of caecilians based on DNA sequence data
was published. Hedges et al. (1993) analyzed partial 16S and 12S mt rDNA
sequences for 13 caecilian species in 9 genera, including members of four of
the family-level taxa recognized in our 1989 classification. Recently, the
taxonomic coverage for these molecular markers has begun to expand, so
that comparative sequence data are now available for 23 species, 16 genera
1
Department of Zoology, The Natural History Museum, London SW7 5BD, United Kingdom
2
Division of Amphibians and Reptiles, Museum of Zoology, The University of Michigan, Ann
Arbor, Michigan, 48109-1079, USA
*
This work is dedicated to the memories of John Eric Wilkinson and Annie Wilkinson.
40 Reproductive Biology and Phylogeny of Gymnophiona
and for representatives of all six families (Wilkinson et al. 2002; 2003b).
Gower et al. (2002), in a study focusing on ichthyophiid caecilians,
demonstrated the potential for sequence data from ribosomal and protein
coding (cytochrome B) mt DNA to help resolve low level taxonomic
problems when the taxonomic sampling is sufficiently dense, and Gower et
al. (2005) tentatively identified an undescribed cryptic species of Sri Lankan
Ichthyophis using molecular data. Recently, San Mauro et al. (2004)
addressed relationships among single representatives of each of the six
families with a combination of complete mitochondrial genomes and RAG-
1 nuclear gene sequences.
Morphological data sets have also been expanded in terms of taxa and
through the discovery of additional characters, and previously assembled
data have been critically reviewed and revised to reduce errors (Naylor and
Nussbaum 1980; Nussbaum and Naylor 1982; Scheltinga et al. 2003, see also
Chapter 7 of this volume; Wilkinson and Nussbaum 1996; Wilkinson 1996,
1997). There have also been a few phylogenetic studies of monophyletic
subgroups at the genus- or species-level using morphological data
(Nussbaum and Hinkel 1994; Wilkinson and Nussbaum 1999; Wilkinson et
al. 2004).
In our 1989 classification, we provided diagnoses of caeciliid genera
based on a core set of characters. Although each genus was understood to
have a unique combination of characters, uniquely derived characters
supporting the monophyly of most caeciliid genera were simply unknown,
and knowledge of the diversity within the more speciose nominate genera
was limited to one or a few species. Since 1989, a single genus and 16
species (one of which we consider invalid) have been newly described, and
5 species have been removed from synonymy. In the same period, two
genera and 5 species have been lost to synonymy (in addition to those we
excluded from our treatment and subsequently synonymized). Little else
has changed, and the limited low-level taxonomic activity belies the fact
that taxonomy at the species- and genus-level remains in need of careful
study and stabilisation. The new genus, Atretochoana was established to
receive a single species of typhlonectid caecilian with a radically divergent
morphology discovered in the course of routine taxonomic work
(Nussbaum and Wilkinson 1995). Atretochoana is the largest lungless
tetrapod and the only known lungless caecilian, and it possesses many
unique features associated with a novel cranial architecture (Wilkinson and
Nussbaum 1997). Its discovery represents a substantial increase in the
perceived diversity of caecilians, and of tetrapods (Donoghue and Alverson
2000), and it serves to emphasise the limited knowledge of caecilian
biodiversity.
Overall, taxonomic coverage has remained patchy in both
morphological and molecular phylogenetic studies. Consequently, even
where inferred relationships for the subset of sampled taxa are well-
supported, they are not readily translated into a phylogenetic classification
of the entire Order. This is exacerbated by the low-level taxonomic
Caecilian Phylogeny and Classification 41
uncertainties that are currently a major obstacle to progress in caecilian

systematics and for caecilian biology more generally. However, some
phylogenetic relationships have been confidently established, some more
tentatively so, and, importantly, we have a clearer picture of what remains
to be done. In this chapter, we present an overview of current
understanding of caecilian phylogeny and an update of our 1989
classification.
2.2 CAECILIAN PHYLOGENY

Caecilians constitute one of three extant orders of the amphibian subclass
Lissamphibia, which includes all of the extant Amphibia. The caecilians,
frogs and toads, salamanders and newts are generally believed to comprise
a monophyletic group based on a variety of presumed shared, derived
character states such as smooth (externally scaleless) epidermis and the
presence of gonadal fat bodies (e.g., Parsons and Williams 1963). That
Lissamphibia is monophyletic with respect to extant taxa is strongly
supported by molecular data which also tend to support either a sister
group relationship between caecilians and salamanders or between
caecilians and salamanders plus frogs (e.g., Feller and Hedges 1998; San
Mauro et al. 2004). However, various hypotheses of polyphyletic origins of
the three orders from different fossil taxa have been proposed, and
monophyly with respect to several extinct groups of Amphibia is far from
uniformly accepted (e.g., Milner 1993; Schoch and Milner 2004).
Regardless of origins and relationships, the three extant orders
(Gymnophiona = caecilians; Caudata = salamanders; Anura = frogs) are
readily distinguished. Many derived characteristics unambiguously
identify all species of Gymnophiona (see Diagnosis below) including the
presence of a unique, dual, jaw-closing mechanism (Nussbaum 1977, 1983;
Fig. 2.1). The presence of paired tentacular sensory organs on the snout on
the edges of, or anterior to, the eyes is unique and can be readily
determined with a hand lens even for very small specimens, although in
some taxa they becomes apparent only at or close to metamorphosis and
they are not present in larvae. Of course, caecilians also have
characteristically elongate, snake-like bodies and completely lack limbs or
girdles.
In 1989 we considered caecilian phylogeny to be poorly understood but
not completely unknown. Nussbaum (1977, 1979, 1985) had provided good
morphological support for the monophyly of two families, the
Rhinatrematidae and Scolecomorphidae, and for two hypotheses of
interfamilial relationships: that the Rhinatrematidae is the sister taxon of all
other caecilians, and that the Scolecomorphidae, Typhlonectidae and
Caeciliidae comprise a monophyletic group. We also accepted the
monophyly of the Typhlonectidae and had a fairly well-supported
phylogeny for the four then recognised typhlonectid genera (Wilkinson
1989). We considered the caeciliids of the Seychelles archipelago to be a
Fig. 2.1. Uniquely derived, dual jaw-closing mechanism, which is diagnostic of caecilians (Nussbaum
1977). The two parts consist of the ancestral jaw-closing mechanism, common to all vertebrates, and
a novel component. In the ancestral mechanism, the m. adductor mandibulae (mam) pulls up on the
lower jaw (d, dentary) in front of the articulation of the lower jaw with the skull. In the novel component,
the m. interhyoideus posterior (mip) pulls down on a process of the dentary (pret, or processus
retroarticularis) that projects posteriorly from the jaw articulation, causing the lower jaw to swing up.
The mip, normally a throat constrictor, takes on a new function of jaw-closing in caecilians. The m.
depressor mandibulae (mdm) serves to open the jaws in all caecilians by pulling up on the pret.
Rhinatrematids have the presumed ancestral condition in which the ancestral jaw-closing mechanism
dominates. Ichthyophiids, caeciliids, and scolecomorphids demonstrate progressively increased
dominance of the novel component. The lower jaw becomes progressively shorter and the pret
progressively longer and more curved dorsally in the same evolutionary sequence. The horizontal bar
= 1 mm. Adapted from Nussbaum, R. A. 1983. Journal of Zoology, London 199: 545-554, Fig. 2.
monophyletic group on the basis of cytogenetic data (Nussbaum and Ducey

1988); and we considered some pairs of caeciliid genera, such as the East
African Afrocaecilia and Boulengerula, Central American Dermophis and
Gymnopis, and South American Caecilia and Oscaecilia, to be obviously
closely related.
In this section we review current understanding of caecilian phylogeny.
Where possible we identify uniquely derived features that unambiguously
support hypothesised monophyletic groups of caecilians. Note that inferred
relationships of families and of genera that are not monotypic are mostly
based on observations of only a subset of the constituent species. Detailed
morphological observations and molecular data are lacking for the vast
majority of caecilian species. Thus, where we list uniquely derived features
supporting the monophyly of a particular group, we do so with the strong
caveat that these features may be unknown for some or most of the
included species. The relationships we discuss are summarised in Figure 2.2
2.2.1 Rhinatrematidae
Nussbaum (1977) listed six uniquely derived features that support the
monophyly of the Rhinatrematidae, a small Neotropical family including
the two genera Rhinatrema and Epicrionops and nine recognised species.
Most distinctive of the supporting conditions is the presence of a posterior
notch in the squamosal that accommodates a distinct process of the os
basale. Apart from the lack of a distinct basipterygoid process, Nussbaum’s
(1977) other derived features of rhinatrematids, the reduction or absence of
ceratobranchials 2 and 3; the larynx posterior to the glossal skeleton; and
the absence of the musculus subarcualis rectus II and III, all relate to the
reduction of the posterior hyobranchial apparatus and are unlikely to be
completely independent. A musculus subarcualis rectus II is now known to be
absent in typhlonectids also, although in typhlonectids its absence is
associated with the elaboration, rather than reduction, of the
buccopharyngeal pump (Wilkinson and Nussbaum 1997), and is surely
convergent with the condition in rhinatrematids. Wilkinson (1996)
identified two derived cardiovascular features that also support
rhinatrematid monophyly, namely the partial division of the normally
undivided sinuatrial aperture and the left pulmonary artery supplying the
oesophagus rather than the left lung.
None of these features has been documented in all the currently
recognised rhinatrematid species, most of which remain unstudied in any
detail, but most have been documented for the monotypic Rhinatrema and
for one or more species of Epicrionops. Based on the general similarity of all
known rhinatrematids, we do not expect these morphological features to
vary much within the family. Rhinatrematid sampling in Wilkinson’s (1997)
morphological phylogenetic analysis and in the molecular study of Gower
et al. (2002) was limited to only two species of Epicrionops in the former and
one Epicrionops and the monotypic Rhinatrema in the latter. In both analyses,
the results were consistent with rhinatrematid monophyly, and the
separation of the rhinatrematids from all other caecilians was well

supported. Phylogenetic relationships within the Rhinatrematidae are as
yet unstudied. Epicrionops differ from Rhinatrema primarily in having a
longer tail with well-developed haemal arches, a more longitudinal vent,
and in retaining a small ceratobranchial 3, all of which are probably
ancestral character states. Derived features that support the monophyly of
Epicrionops are currently unknown.
2.2.2 Non-rhinatrematids — Neocaecilia

The discredited phylogenies of Laurent (1986) and Lescure et al. (1986)
notwithstanding, since Nussbaum’s (1977, 1979) pioneering morphological
phylogenetic studies, the basal split in the caecilian tree has been thought
to be between the rhinatrematids and all other caecilians. Nussbaum (1977)
identified 13 morphological features of rhinatrematids that he considered
ancestral and unique within caecilians, with the corresponding derived
character states supporting the monophyly of all non-rhinatrematid
caecilians. Of these, we currently view five as uniquely derived: sides of
parasphenoid converge anteriorly; no contact between quadrate and
maxillopalatine; long and recurved retroarticular processes; musculus
interhyoideus posterior has no insertion on the ceratohyal; and musculi
adductores mandibulae externi do not meet mid-dorsally. The presence of two
rows of trunk neuromasts in larvae is interpreted as an additional unique
and ancestral feature of rhinatrematids (Wilkinson 1992a), and Wilkinson
(1996) identified five uniquely derived cardiovascular features that also
support the monophyly of the non-rhinatrematids: ventricle relatively
narrow and elongate; a bipartite sinus venosus; conus arteriosus not bent to
the left; an elongate truncus arteriosus; and common systemicocarotid
arteries.
Until recently, molecular data have been mostly consistent with, but
had not provided compelling support for, the basal split between
rhinatrematids and all other caecilians (Hedges et al. 1993). Most recent
analyses, both morphological (Wilkinson and Nussbaum 1996; Wilkinson
1997) and molecular (Gower et al. 2002; Wilkinson et al. 2002, 2003b) have
assumed that the Rhinatrematidae is the sister group of all other caecilians
in order to root the caecilian tree and have not provided any further test of
this hypothesis. Although taxon sampling is limited, the extensive
mitochondrial and nuclear gene data of San Mauro et al. (2004) provide very
strong support for non-rhinatrematid monophyly, and we consider it to be
a very well supported hypothesis on the basis of diverse morphological and
molecular data.
Canatella and Hillis (1993) coined the term Stegokrotaphia for the clade
including all non-rhinatrematids. However, this name belies a great
diversity of zygokrotaphic scolecomorphids, caeciliids and typhlonectids
within the group. For this reason we prefer the anatomically neutral
Neocaecilia as a rankless epithet for the suprafamilial clade including all
caecilians with jaw-closing muscles that do not extend onto the top of the
skull from the adductor chamber.
2.2.3 Ichthyophiidae
The Ichthyophiidae comprises some 39 species in the two genera
Caudacaecilia and Ichthyophis. Caudacaecilia is restricted to South East Asia
whereas Ichthyophis also has representatives in South Asia. In Nussbaum’s
(1979) analyses, Caudacaecilia and Ichthyophiidae were closely related, but
lacked any uniquely derived features that supported their monophyly. In
fact, despite the external similarity of all ichthyophiids, and practical
difficulty of distinguishing ichthyophiid species (Nussbaum and Gans
1980), only a single feature that supports ichthyophiid monophyly has been
reported previously, the presence of angulate annuli on the anteroventral
surface (Nussbaum 1977; Wilkinson and Nussbaum 1996). An additional
supporting derived feature, a short parasphenoid that does not extend as
far anteriorly as the posterior margin of the choanae, is characteristic of all
ichthyophiid skulls and of no non-ichthyophiid skulls that we have
examined.
Wilkinson (1997) and Wilkinson et al. (2002) both recovered the very
few ichthyophiids included in their morphological and molecular
phylogenetic analyses (three and two species of Ichthyophis respectively) as
a reasonably well-supported monophyletic group. In contrast, the
molecular study of Gower et al. (2002) included a broader range of
Ichthyophis species and yielded optimal trees in which the Ichthyophiidae is
paraphyletic with respect to the Uraeotyphlidae. Although this result was
not significantly better supported than alternative trees in which the
Ichthyophiidae is monophyletic, and thus does not justify any taxonomic
changes at this time, it raises the possibility of ichthyophiid paraphyly and
indicates the need for further phylogenetic study and additional character
data. Phylogenetic relationships among ichthyophiids are poorly
understood, but Gower et al. (2002) suggested that the Ichthyophis of Sri
Lanka, and those of South East Asia comprise distinct monophyletic groups
(see also Bossuyt et al. 2004; Gower et al. 2005). All recent work supports the
idea that the ichthyophiids were present on the Indian plate prior to its
collision with Laurasia, and that South East Asian ichthyophiids result from
one or more dispersals out of India (Gower et al. 2002; Wilkinson et al. 2002).
Caudacaecilia and Ichthyophis are differentiated by the absence or
presence of splenial teeth in adults respectively. Absence of splenial teeth is
considered derived within caecilians (Nussbaum 1979) but appears to be
quite homoplastic (Nussbaum and Wilkinson 1989). Thus there is only very
weak evidence known to support the monophyly of Caudacaecilia, no
known derived features supporting the monophyly of Ichthyophis, and a
strong possibility that Ichthyophis is paraphyletic with respect to
Caudacaecilia. No Caudacaecilia have been included in any numerical
phylogenetic analysis since Nussbaum (1979).
2.2.4 Uraeotyphlidae
The genus Uraeotyphlus comprises five nominate species from southern
peninsular India. The genus was included in the Caeciliidae by Taylor
(1968, 1969) but transferred to its own subfamily within the Ichthyophiidae
by Nussbaum (1979). Duellman and Trueb (1986) elevated Nussbaum’s
Uraeotyphlinae to family level because their phylogenetic analyses
suggested that inclusion of the Uraeotyphlinae within Ichthyophiidae
rendered the latter paraphyletic. Uraeotyphlids share a combination of
ancestral and derived features, but there are no known uniquely derived
features that support the monophyly of the genus and family, a
consequence of extensive convergence between uraeotyphlids and various
caeciliids. Gower et al. (2002) included three uraeotyphlid species in their
molecular phylogenetic analysis and obtained strong support for
monophyly of the group as a whole as well as for relationships among the
species. The anterior tentacles, dorsal nares, and recessed subterminal
mouths, though not unique, are probably derived within the
Ichthyophiidae plus Uraeotyphlidae, providing qualified support for the
monophyly of Uraeotyphlus (Wilkinson and Nussbaum 1996).
2.2.5 Ichthyophiidae + Uraeotyphlidae — Diatriata

Nussbaum (1979), and subsequently both Duellman and Trueb (1986) and
Hillis (1991), recovered the Uraeotyphlidae as more closely related to
caeciliids, scolecomorphids and typhlonectids than to ichthyophiids. The
major change in our understanding of higher caecilian phylogeny has been
the adoption of the alternative hypotheses that the Uraeotyphlidae plus
Ichthyophiidae are a clade. This hypothesis was well supported by
morphological data that incorporated newly discovered cardiovascular
characters (Wilkinson and Nussbaum 1996; Wilkinson 1997), and it has
subsequently received strong support from analyses of gene sequence data
(Wilkinson et al. 2002, 2003b; San Mauro et al. 2004). We currently interpret
five uniquely derived features to support the Uraeotyphlidae +
Ichthyophiidae clade: circumorbital bone (often termed postfrontal)
present, external division of the atrium, an elongate anterior pericardial
space, two posterior internal flexures in the musculus rectus lateralis, and an
internal flexure in the musculus subvertebralis. The first of these is somewhat
variable in that it may be more or less fused to adjacent elements, and it has
been previously considered primitive, but, at least within caecilians, the
reverse polarity is more parsimonious. We find the strong and congruent
support from morphology and from molecules for this phylogenetic
hypothesis to be compelling. We propose Diatriata, as a suitable rankless
name for this suprafamilial clade comprising those caecilians with partial
external division of their atrium.
2.2.6 The Higher Caecilians — Teresomata

The sister group of the Uraeotyphlidae + Ichthyophiidae, is a clade
comprising the Scolecomorphidae, Typhlonectidae and the Caeciliidae, and
informally termed the advanced (Nussbaum 1991) or higher (e.g., San

Mauro et al. 2004) caecilians. We know of only two putatively unique and
derived features of the advanced caecilians, the absence of a true tail, with
a true tail being defined compositely as a tapering postcloacal region with
internal (vertebrae) and external segmentation (annuli), and the absence of
internal flexures in the musculus rectus lateralis (Nussbaum and Naylor 1982;
Wilkinson 1997). However, this grouping has been recovered in all
numerical phylogenetic analyses of morphology or molecules, and with
strong support. Compared to its sister clade, the advanced caecilians appear
to be more speciose, and the group is much more diverse in morphology,
ecology and life history. Its distribution is more cosmopolitan, with
representatives in all areas where caecilians are found except South East
Asia. The informally named ‘advanced caecilians’ has stood the test of time,
and we suggest Teresomata as a rankless name for this suprafamilial clade,
which encompasses the Scolecomorphidae, Typhlonectidae, and
paraphyletic Caeciliidae. These caecilians lack true tails and have, for the
most part, more rounded (teres) ends to their bodies (soma) than
rhinatrematids, ichthyophiids and uraeotyphlids.
2.2.7 Scolecomorphidae
The Scolecomorphidae was established by Taylor (1969) for a few
distinctive African caecilians. It currently comprises the West African
Crotaphatrema and East African Scolecomorphus, each with three nominate
species. These genera share many distinctive, derived morphological
features that provide strong support for scolecomorphid monophyly. These
include the absence of stapes and foramina ovales, absence of internal
processes on lower jaws, a transverse bar extending between the
posteromedial edges of the posteriormost ceratobranchial elements of the
glossal skeleton (Nussbaum 1977), and a mobile eye attached to the base of
the tentacle (Taylor 1968; Nussbaum 1981, 1985; O’Reilly et al. 1996). No
Crotaphatrema have been included in any numerical phylogenetic analyses,
but Wilkinson (1997) included all three Scolecomorphus species in his
morphological phylogenetic analysis, and two species were included in the
Wilkinson et al. (2003b) molecular phylogenetic study. In both cases there
was strong support for the monophyly of Scolecomorphus, which is
supported by the presence of a uniquely large diastema between the
vomerine and palatine dental series. Monophyly of Crotaphatrema is
supported by the particular form of stegokrotaphy in which the upper
temporal fossa is obliterated by an outgrowth of the parietal (Nussbaum
1985). Wake (1998) suggested a close relationship between S. kirkii and S.
vittatus on the basis of similar phallus morphology.
2.2.8 Typhlonectidae
The Typhlonectidae was established by Taylor (1968) for a group of
Neotropical caecilians that he believed were aquatic and which are now
considered to be either aquatic or semi-aquatic (Nussbaum and Wilkinson
1987; 1989). As currently conceived, the family includes five genera, three
of which are monotypic. A highly distinctive derived feature that supports
typhlonectid monophyly is the fused, sac-like form of the foetal gills
(Wilkinson 1989; Wilkinson and Nussbaum 1999). This feature is known in
all typhlonectids for which fetuses have been examined but remains
unknown in Potomotyphlus and Atretochoana. Wilkinson and Nussbaum
(1999) identified six additional features that appeared to be unique and
derived in typhlonectids. These are: small tentacular apertures and (non-
protrusible) tentacles, relatively dorsally oriented occipital condyles, a
ventral process of the squamosal bracing against the maxillopalatine, M-
shaped ceratohyals, a sliding articulation between the third and fourth
ceratobranchials, and the musculus subvertebralis pars ventralis with a
scalloped origin.
Wilkinson’s (1997) morphological phylogenetic analysis found strong
support for the pairing of the only two typhlonectids, Chthonerpeton
indistinctum and Typhlonectes natans, that it included, and is the only study
to have provided a numerical phylogenetic test of typhlonectid monophyly
(which has never been seriously questioned). Well-supported relationships
within the Typhlonectidae have been inferred on the basis of extensive
morphological data (Wilkinson and Nussbaum 1999; see Fig. 2.2). Although
monophyly of Chthonerpeton has not been established, the generotype, C.
indistinctum, lies outside a group including all other typhlonectid genera.
Nectocaecilia, which is believed to be semi-aquatic, is the sister group of a
clade of fully aquatic, finned caecilians that comprises Atretochoana,
Potomotyphlus, and Typhlonectes. Potomotyphlus appears to be the sister
genus of the lungless Atretochoana on the basis of a number of features
associated with a reduction in pulmonary respiration.
2.2.9 Caeciliidae
As currently conceived, the Caeciliidae appears to be a relatively
heterogeneous and paraphyletic assemblage comprising all those caecilians
that have never been removed to another family. Molecular data strongly
support the paraphyly of the Caeciliidae with respect to the
Typhlonectidae, recovering Caecilia as more closely related to Typhlonectes
than to a broad range of other caeciliids (Hedges et al. 1993; Wilkinson et al.
2002, 2003b). Hedges et al. (1993) proposed removing caeciliid paraphyly by
recognising the typhlonectids at the sub-familial rather than familial level,
but in the absence of a better understanding of the relationships among
higher caecilians this action would only shift the problem of paraphyly to
a different taxonomic level. Paraphyly of the Caeciliidae with respect to the
Scolecomorphidae is also suggested by the most recent molecular
phylogenetic study (Wilkinson et al. 2003b). Phylogenetic analyses based on
morphology are less clear cut, with caeciliid monophyly or paraphyly with
respect to both the Typhlonectidae and Scolecomorphidae achieved under
alternative weighting schemes (Wilkinson 1997) and thus not well-
supported. A single derived feature, an elongate musculus interhyoideus
posterior (Fig. 2.1), supports the monophyly of the Caeciliidae. However,

given the strong molecular support for caeciliid paraphyly, the shorter form
of the muscle in typhlonectids must be presumed to be due to reversal,
presumably associated with their zygokrotaphy.
We have only limited understanding of the relationships among
caeciliids. Some relationships are suggested by taxonomic history. Oscaecilia
was established by Taylor (1968) through the partitioning of Caecilia on the
basis of a single difference (the eye covered with bone or not) of dubious
value. Although both genera are thus of uncertain monophyly, we are
content to assume that they are jointly monophyletic based on their overall
similarity in external morphology and cranial architecture. Similarly,
Luetkenotyphlus was established through partitioning of Siphonops, and these
genera are presumed to be closely related to each other and to
Mimosiphonops (Wilkinson and Nussbaum 1992), although we know of no
uniquely derived characters of this group or of any of its constituent
genera. The Central American species of the genera Dermophis and
Gymnopis are presumed to be jointly monophyletic on the basis of their
overall similarity in morphology and reproduction. They differ in the
presence or absence of splenial teeth and whether the eye is covered with
bone, features which appear highly homoplastic in caecilians and which do
not convince us of the monophyly of either genus. Of the caeciliid taxa
studied thus far, molecular data strongly support a close relationship of
Dermophis (and by implication Gymnopis) with the African genus
Schistometopum, reflecting a previous taxonomic association.
Among Old World caeciliids, molecular evidence supports the
monophyly of the caeciliids of the Seychelles, which Nussbaum and Ducey
(1988) had argued previously on the basis of cytological data. All species of
the Seychellean caeciliid clade have been included in molecular
phylogenetic analyses, an atypical level of taxonomic coverage for
caecilians. These data convincingly identify Praslinia as the sister taxon to
a Grandisonia + Hypogeophis clade, without resolving the relationships
within this latter clade. They suggest that Grandisonia, which (like the
Seychellean clade as a whole) lacks any known uniquely phenotypic
derived traits, is paraphyletic with respect to Hypogeophis. Among the
sampled taxa, the molecular data also provide strong support for the Indian
caeciliid Gegeneophis being the sister-group of the Seychelles caeciliids
(Wilkinson et al., 2002; 2003b), suggesting the possibility of an Indo-
Seychellean caeciliid clade that we might expect to also include the thus far
unstudied Indian caeciliid Indotyphlus. An Indo-Seychellean connection for
caecilians was predicted on the basis of biogeographic and plate tectonics
considerations (Nussbaum 1984). Molecular data also support, albeit not
strongly, the pairing of the East African Boulengerula and West African
Herpele.
Wilkinson (1997) considered the available morphological data to be
insufficient to unravel relationships within the higher caecilians. We
interpret the failure of analyses based on morphology to support those
relationships that are well supported by molecular data, including the

monophyly of the Seychellean caeciliids, the Indo-Seychellean clade, and
the Dermophis-Schistometopum grouping, to further indicate the current
limitations of the morphological data rather than undermining the
molecular results. Morphological data have provided very useful
phylogenetic characters, but broader and deeper sampling of characters and
taxa are needed, particularly within the higher caecilians. Although
relationships within the higher caecilians are poorly resolved in our
relatively conservative consensus phylogeny (Fig. 2.2), sampling of taxa for
both morphological and molecular phylogenetic study is improving.
2.2.10 Prospects
The relatively small number of species of Gymnophiona means that a fairly
comprehensive phylogeny for the major lineages of caecilians is a realistic
Fig. 2.2. Summary (consensus) phylogeny of caecilian constructed manually and based on the results
of previous numerical phylogenetic analyses and inferences from taxonomy as described in the text.
The dashed line indicates particularly uncertain monophyly highlighting the potential for the branches
from the main polytomy, including many unstudied taxa, to perhaps lie within this group. Note that
monophyly of many genera is at best uncertain and that there is a need for much more detailed, low-
level, taxonomic work. Original.
short-term goal that we believe is within reach, and will be reached in the
near future. At lower taxonomic levels the systematic foundations are not
so good. Until about 1972, caecilian taxonomy was dominated by E. H.
Taylor who described many species that have not withstood subsequent
scrutiny and have been lost to synonymy. The reduction in numbers of
recognised species has more or less balanced the description of new species
since Taylor’s time, and the work of checking and testing Taylor’s species-
level taxonomic work remains largely incomplete. Most caecilian species
are poorly known and poorly circumscribed, impinging on all aspects of
their biology (e.g., Gower and Wilkinson, 2005), and taxonomic uncertainty
and instability can be expected to continue for some time. However,
molecular phylogenetic studies at low taxonomic levels are proving useful
in helping to delimit and distinguish morphologically similar species in
genera such as Ichthyophis, where identification to species is notoriously
difficult and the existing taxonomy exceedingly problematic. Molecular
studies should facilitate taxonomic work while providing the low-level
phylogenies needed to study caecilian evolution in detail. Ultimately any
understanding of caecilian species, be it derived from molecules or
morphology or (preferably) both, will depend on collecting sufficient (i.e.,
much more) new material across many taxa. Traditionally, caecilians have
been poorly sampled in the field, but this is improving. We think it likely
that many new caecilian species will be discovered, both in research
collections and through additional collecting. It is noteworthy that the
majority of new species described in the 15 years since our last review have
been described in the last five years, perhaps indicating a rise in interest
and the beginnings of a new era of taxonomic discovery (e.g., Gower et al.,
2004). As caecilian taxonomy stabilises, and the rate of losses to synonymy
decreases, we expect the overall number of recognised caecilian species to
increase, and we suspect that the number of currently recognised species is
a considerable underestimate of their actual diversity.
2.3 CLASSIFICATION
The classification presented here includes no major changes from our
previous summary (Nussbaum and Wilkinson 1989), because we think
none is warranted by the current state of knowledge. We draw attention to
three important studies of the taxonomy of regional caecilian faunas by
Pillai and Ravichandran (1999), Savage and Wake (2001) and Lynch (1999)
that provide recent keys. We prefer to accept the paraphyly of the
Caeciliidae for the time being, in the belief that the best way of removing
it will only become apparent with a more comprehensive understanding of
the relationships of caeciliid genera, particularly those that remain
unstudied phylogenetically. Formal taxonomic revision aimed at removing
the paraphyly now, although well-intended, would be incomplete and
unlikely to promote stability in the meanings of names. Thus we use the
same six-family system and same format as in 1989, but with updated
species lists and synonymies. Where possible, we have indicated derived

features for genera that are not monotypic and where this has not been
discussed above, but our diagnoses rely upon unique combinations of
features, not all of which are themselves unique. Reference to a
pseudoectopterygoid (Wilkinson and Nussbaum 1992) is to the separate
palatal bone that lies between the pterygoid (process of the quadrate) and
the maxillopalatine in some caeciliids. We also prefer to use inner
mandibular for those teeth usually referred to as splenials. In both cases our
usage is intended to avoid asserting homologies for which there is no good
evidence.
ORDER GYMNOPHIONA RAFINESQUE 1814

Diagnosis. Lissamphibia without limbs and girdles; with paired sensory
tentacles on the snout; with a dual jaw-closing mechanism consisting of the
ancestral component (musculus adductor mandibulae pulling up on ramus of
lower jaw) and a unique, novel component (musculus interhyoideus posterior
pulling back and down on retroarticular process of lower jaw); with an
eversible phallus in males formed by the posterior part of the cloaca; with
an os basale; trunk vertebrae with enlarged basipophyseal processes; atlas
without a tuberculum interglenoideum.
Content. 6 families, 33 genera, 170 species.
Distribution. Pantropical, except for Madagascar and southeast of
Wallace’s Line. Distribution includes some subtropical areas.
Remarks. Jenkins and Walsh (1993) described a ‘caecilian’ fossil, Eocaecilia,
from the Jurassic of North America and more recently Evans and
Sigogneau-Russell (2001) described a ‘stem-group caecilian’ Rubricacaecilia
from the Lower Cretaceous of North Africa. This fossil taxon, based on a
brief description that has yet to be significantly expanded, has seemingly
been universally accepted as a caecilian with little, if any, critical discussion.
Unlike living caecilians, Eocaecilia has small limbs, and thus, by the
diagnosis given above, it is not a gymnophionan (= caecilian). The presence
or absence of limbs in the fragmentary Rubricacaecilia is unclear, but it has
other features, notably the atlas has a tuberculum interglenoideum, which
places it outside our Gymnophiona. Evans and Sigogneau-Russell (2001)
supported Trueb and Cloutier’s (1991) use of “Apoda Oppel 1811 for the
crown group alone and Gymnophiona Rafinesque for the clade comprising
stem-group taxa + Apoda”, a use also adopted by Schoch and Milner
(2004). Our previous and present diagnoses of Gymnophiona are based on
the living species, and they exclude what Evans and Sigogneau-Russell
view as stem-group caecilians. Neontologists use the term Gymnophiona to
convey generalities about the living caecilians that may not hold or may not
be known in the stem group. Changing the meaning of Gymnophiona
means that general statements in the literature, such as ‘Gymnophiona are
legless’, become technically incorrect, and neontologists, who comprise the
bulk of caecilian researchers would have to learn to adopt Apoda in order
to make generalisations about the living caecilians. The latter name is also
problematic because zoological classification is festooned with other uses of
it, most importantly the homonymy with Apoda Haworth 1809, a genus of
moth. We believe clarity would be best served if the use of Apoda in
caecilian classification were completely abandoned. We see no good reason
to rediagnose Gymnophiona in order to accommodate relatively poorly
known fossil taxa that, while possibly closely related to caecilians, are not
caecilians in the sense in which the term is generally used. A. Family
Rhinatrematidae Nussbaum 1977 (Fig. 2.3 A, B)
Diagnosis. Gymnophiona with true tails consisting of a postcloacal
segment with vertebrae, myomeres, and complete skin annuli; primary
annuli divided by secondary and tertiary grooves; all annular grooves
orthoplicate; numerous scales in all annular grooves and in some of the
dorsal grooves of the collars; strongly zygokrotaphic skulls with the musculi
adductores mandibulae externi passing through the temporal fossae to meet at
the midline of the skull along the interparietal suture; maxillopalatine in
contact with the quadrate; squamosal widely separated from the frontal,
notched posteriorly, the notch opposing a dorsolateral process of the os
basale; premaxillae and nasals present as separate bones; mouth terminal;
retroarticular process of lower jaw short and not curved dorsally; musculus
interhyoideus posterior short; stapes pierced by stapedial artery; tentacle
immediately anterior to or on the anterior edge of eye; eyes visible
externally, in a socket in the maxillopalatine; hyobranchium of adults with
only three ceratobranchial elements decreasing in size posteriorly, with the
larynx situated posterior to the hyobranchium (not enclosed between the
two arms of the posteriormost ceratobranchials); hyobranchial elements of
larvae mineralized, hyobranchium of metamorphosed individuals
cartilaginous; truncus arteriosus short; atrium undivided externally.
Content. 2 genera, 9 species.
Distribution. Northern South America.
Remarks. There has been no change in the taxonomy of the
Rhinatrematidae since our review of 1989, and no species description or
other taxonomic actions since Taylor (1968).
1. Epicrionops Boulenger 1883
Type species: Epicrionops bicolor by original monotypy.
Diagnosis. Rhinatrematids with three ceratobranchial arches in adults; a
longitudinal cloacal opening; relatively long tail consisting of more than 11
postcloacal annuli; more than one row of scales per annular groove.
Content. 8 species: bicolor, columbianus, lativittatus, marmoratus, niger, parkeri,
peruvianus, petersi.
Distribution. Colombia, Ecuador, Peru, and Venezuela.
2. Rhinatrema Duméril & Bibron,1841
Type species: Caecilia bivittata Guérin-Méneville 1829, by monotypy.
Colour
Figure
Fig. 2.3 contd

Diagnosis. Rhinatrematids with two ceratobranchial arches in adults; more

transverse or subcircular cloacal opening; relatively short tail consisting of
11 or fewer postcloacal annuli; a single row of scales per annular groove.
Content. l species: bivittatum.
Distribution. Brazil, French Guiana, Guyana, and Surinam.
Remarks. Nussbaum and Hoogmoed (1979) noted the rarity of Rhinatrema
bivittatum in scientific collections. Recent fieldwork in French Guiana
yielded considerable additional material suggesting the species is not
particularly rare in the wild (Wilkinson, Gower and Kupfer, unpublished).
B. Family Ichthyophiidae Taylor 1968 (Figs. 2.3C, D; 2.6A, B)
Diagnosis. Gymnophiona with true tails; skull stegokrotaphic; musculi
adductores mandibulae externi confined beneath the skull roof, not meeting
middorsally; distinct septomaxillae, premaxillae, nasals, and prefrontals;
circumorbitals (postfrontals) distinct or partially or entirely fused to
maxillopalatine or squamosal; frontal and squamosal in contact; no
dorsolateral process on os basale; no posterior notch in squamosal; quadrate
and maxillopalatine broadly separated; stapes pierced by stapedial artery;
mouth nearly terminal; retroarticular process of lower jaw curved dorsally;
musculus interhyoideus posterior short; tentacular opening between the eye
and nostril, usually closer to the eye and below the eye-nostril line;
ceratohyal arch U-shaped; four ceratobranchial arches in larvae, arches 3
and 4 fused in adults; larynx positioned between the distal ends of fused
arches 3 and 4; all primary annuli subdivided by secondary and tertiary
grooves in metamorphosed individuals; annular grooves angulate ventrally
over most of the body, orthoplicate posteriorly only; numerous scales
present in all but perhaps a few anterior annular grooves of adults; aortic
arches proximal to the heart fused into an elongate truncus arteriosus; atrium
partially divided externally.
Distribution. India, Sri Lanka, and southeast Asia including southern
Philippines and Indo-Malaysian Archipelago northwest of Wallace’s Line.
1. Caudacaecilia Taylor 1968
Type species. Ichthyophis nigroflavus Taylor 1960, by original designation.
Diagnosis. Ichthyophiids without inner mandibular teeth.
Content. 5 species: asplenia, larutensis, nigroflava, paucidentula, weberi.
Distribution. Borneo, Malay Peninsular, Philippines, Sumatra.
Remarks. Taylor (1968) and Nussbaum and Gans (1980) examined
specimens of Caudacaecilia (cf. asplenia) from Sri Lanka in museums, but
Fig. 2.3 contd
Fig. 2.3 A-B. Adult Rhinatrema bivittatum (Rhinatrematidae) from French Guiana; the tentacle (small
white spot) is on the anterior edge of the eye; photo by Peter Stafford. C-D. Adult Ichthyophis
bannanicus (Ichthyophiidae) from Mengla, Yunnan, China; the tentacle is just above the mouth and
closer to the eye than to the naris; photo by Edmund D. Brodie, Jr.
concluded that there was some doubt about the collection data. Recent
caecilian surveys in Sri Lanka have not revealed additional specimens
attributable to this genus.
2. Ichthyophis Fitzinger 1826
Type species. Caecilia glutinosa Linnaeus 1758, by original monotypy.
Diagnosis. Ichthyophiids with splenial teeth.
Content. 34 species: acuminatus, atricollaris, bannanicus, beddomei,
bernisi, biangularis, billitonensis, bombayensis, dulitensis, elongatus,
garoensis, glandulosus, glutinosus, humphreyi, husaini, hypocyaneus,
javanicus, kohtaoensis, laosensis, longicephalus, malabaricus,
mindanaoensis, monochrous, orthoplicatus, paucisulcus, peninsularis,
pseudangularis, sikkimensis, singaporensis, subterrestris, sumatranus,
supachaii, tricolor, youngorum.
Distribution. South East Asia, India, Sri Lanka, southern Philippines,
western Indo-Australian Archipelago.
Remarks. Two new species of Ichthyophis (husaini and garoensis) were
described by Pillai and Ravichandran (1999), and Gower et al. (2005)
suggested the presence of an undescribed cryptic species in Sri Lanka.
Kupfer and Müller (2004) provided a rediagnosis of I. supachii. I.
longicephalus (Pillai 1986) was overlooked and not included in our previous
treatment (Nussbaum and Wilkinson 1989).
C. Family Uraeotyphlidae Nussbaum 1979 (Fig. 2.4 A, B)
Diagnosis. Gymnophiona with true tails; weakly stegokrotaphic skulls; m.
adductor mandibulae externi confined beneath the skull roof but may be
visible through a small opening between the squamosal and parietal;
number and arrangement of skull and lower jaw bones and configuration
of the hyobranchium as in the Ichthyophiidae; Stapes imperforate; m.
interhyoideus posterior short; mouth recessed or subterminal; tentacular
opening far forward, below nostril; external nares relatively dorsal, most
primary annuli divided by secondary grooves, a few anterior primary
annuli may not be subdivided, or primary and higher-order annuli
indistinguishable externally; annular grooves do not completely encircle the
body; scales present; aortic arches proximal to the heart fused into an
elongate truncus arteriosus; atrium partially divided externally.
Content. l genus, 5 species.
Distribution. Southern peninsular India.
Remarks. The diagnosis (of Nussbaum and Wilkinson 1989) has been
modified to account for new information on the diversity of uraeotyphlid
annulation patterns (Gower and Wilkinson in prep.; Nussbaum, pers. obs.).
1. Uraeotyphlus Peters 1879
Type species. Coecilia oxyura Duméril and Bibron 1841, by subsequent
designation of Noble (1924).
Diagnosis. As for the family.
Content. 5 species: interruptus, malabaricus, menoni, narayani, oxyurus.

Distribution. Southern peninsular India, Kerala, Karnataka and Tamil
Nadu.
Remarks. Pillai and Ravichandran (1999) described U. interruptus from
Kerala since our (Nussbaum and Wilkinson 1989) review.
D. Family Scolecomorphidae Taylor 1969 (Fig. 2.5 C, D, E)
Diagnosis. Gymnophiona that lack stapes and foramina ovales;
septomaxillae and prefrontals present; no internal process on the
pseudoangular bone; no m. levator quadrati; m. interhyoideus posterior short;
a distinctive hyobranchium in which the flattened distal ends of the fourth
branchial arch are connected by a transverse bar above the larynx; all
primary annuli undivided; aortic arches proximal to the heart fused into an
elongate truncus arteriosus; atrium undivided externally.
Distribution. East and equatorial West Africa.
1. Crotaphatrema Nussbaum 1985
Type species. Herpele bornmuelleri Werner, 1899, by original designation.
Diagnosis. Scolecomorphids without temporal fossae; without diastemata
between the vomerine and palatine series of teeth; and with the maxillary
series extending further posteriorly than the palatine series.
Content. 3 species: bornmuelleri, lamottei, tchabalmbaboensis.
Distribution. Cameroon.
Remarks. Lawson (2000) added C. tchabalmbaboensis to the known species of
Crotaphatrema.
2. Scolecomorphus Boulenger 1883
Type species. Scolecomorphus kirkii Boulenger 1883, by original monotypy.
Diagnosis. Scolecomorphids with temporal fossae; with diastemata
between the vomerine and palatine series of teeth; and with all or most of
the palatine teeth posterior to the maxillary teeth.
Content. 3 species: kirkii, uluguruensis, vittatus.
Distribution. East Africa: Malawi, and Tanzania.
Remarks. The phallus of all Scolecomorphus species is equipped with
cartilaginous spines or spicules (Taylor 1968; Wake 1998). This may be a
uniquely derived character of the genus but the condition in Crotaphatrema
is undocumented.
E. Family Caeciliidae Rafinesque 1814 (Figs. 2.4 C, D; 2.5 A; 2.6 C-F)
Diagnosis. Gymnophiona with nasal and premaxilla fused; septomaxilla,
prefrontal, postfrontal and pterygoid lost or fused to adjacent bones;
pseudoectopterygoid present or not; maxillopalatine widely separated from
quadrate; temporal fossae usually absent, if present, m. adductor mandibulae
externi do not pass dorsally through the fossae; M. interhyoideus posterior
long, with posterior portion extending as far as the sixth trunk myomere;
Colour
Figure
Fig. 2.4 contd

M-shaped ceratohyal arch; larynx between distal ends of fused third and
fourth ceratobranchials; no tail; some, none, or all primary annuli
subdivided by secondary grooves; no tertiary grooves; scales present or
absent; external gills of embryos in three rami (one ramus may be reduced
or vestigial), not fused and sac-like; aortic arches proximal to the heart
fused into an elongate truncus arteriosus; atrium undivided externally.
Distribution. Tropical Central and South America, equatorial East and West
Africa, islands of the Gulf of Guinea, Seychelles Archipelago, and India.
Remarks. During the late 1980’s and early 1990’s, the name “Caeciliaidae”
was used for this family in an attempt to remove the homonymy of
Caeciliidae Rafinesque, 1814 (Amphibia) with Caeciliidae Kolbe, 1880
(Insecta). The problem is outlined in Moore et al. (1984). In 1996, the
Commission on Zoological Nomenclature, under its plenary powers, ruled
(Opinion 1830, BZN 53(1):68-69) that Caeciliidae Rafinesque, 1814 is the
valid amphibian name, and the insect name was changed to Caeciliusidae
Kolbe, 1880.
1. Boulengerula Tornier 1897
Type species. Boulengerula boulengeri Tornier 1897, by monotypy.
Diagnosis. Caeciliids with eye (if present) under bone; no temporal fossae;
mesethmoid exposed between frontals or not; inner mandibular teeth
present or not; no secondary grooves; no scales; tentacular opening nearer
to eye than to external naris; an unsegmented terminal shield; no narial
plugs; a strong diastema between the vomerine and palatine teeth present
or not; a vertical keel on the end of the terminal shield.
Content. 6 species: boulengeri, changamwensis, denhardti, fischeri, taitana,
uluguruensis.
Distribution. Kenya, Malawi, Tanzania and Rwanda.
Remarks. Nussbaum and Hinkel (1994) placed Afrocaecilia in the synonymy
of Boulengerula and described B. fischeri. Wilkinson et al. (2004) removed B.
denhardti from the synonymy of Schistometopum gregorii. Their phylogenetic
analyses using Nussbaum and Hinkel’s morphological data were unable to
resolve well-supported relationships within the genus. Monophyly of the
genus is not seriously in question. Many of the diagnostic features are
derived within the Neocaecilia although none of them uniquely so, and in
molecular analyses, grouping of the two species of Boulengerula included
thus far is well supported (Wilkinson et al. 2003b).
Fig. 2.4 contd
Fig. 2.4 A-B. Adult or subadult Uraeotyphlus sp. (Uraeotyphlidae) from the Western Ghats, peninsular
India; note the dorsal orientation of the eyes and external nares, and the undivided primary annuli on
the anterior portion of the body; photo by John Measey. C-D. Adult Caecilia cf. tentaculata (Caeciliidae)
from South America; note the lack of undivided primary annuli over most of the body, but with some
subdivided primaries posteriorly; the tentacle cannot be seen, because it is directed ventrally from the
“shelf” below the external naris; photo by Peter Stafford.
Colour
Figure
Fig. 2.5 contd

2. Brasilotyphylus Taylor 1968

Type species. Gymnopis braziliensis Dunn 1945, by original designation and
monotypy.
Diagnosis. Caeciliids with eye under bone; no temporal fossae;
mesethmoid covered by frontals; no splenial teeth; secondary grooves
present; scales present; tentacular opening closer to eye than to external
naris; no terminal shield; a very short series of premaxillary-maxillary teeth,
not extending posterior of the choanae; a strong diastema between the
vomerine and palatine teeth; a vertical keel on the body terminus.
Content. l species: braziliensis.
Distribution. Brazil.
Remarks. The affinities of this genus, which has similarities to Boulengerula
in the Old-World and Microcaecilia in the New, are quite unclear at present
(Nussbaum and Hinkel 1994).
3. Caecilia Linnaeus 1758
Type species. Caecilia tentaculata Linnaeus 1758, by subsequent designation
of Dunn (1942).
Diagnosis. Caeciliids with eye not covered with bone; no temporal fossae;
mesethmoid exposed between frontals; splenial teeth present; secondary
grooves present or absent; scales present or absent; subdermal scales
present or absent; tentacular opening directly below external naris, closer to
naris than to eye; unsegmented terminal shield present or not; narial plugs
present; no diastema between vomerine and palatine teeth; no terminal
keel; teeth relatively few and large, usually replaced alternately in groups;
vomeropalatine tooth row displaced posteriorly, not parallel to
premaxillary-maxillary tooth row, diverging from the latter anteriorly
forming an angle where the two rows meet rather than a semicircle.
Content. 33 species: abitaguae, albiventris, antioquiaensis, armata, attenuata,
bokermanni, caribea, corpulenta, crassisquama, degenerata, disossea, dunni,
flavopunctata, gracilis, guntheri, inca, isthmica, leucocephala, marcusi, mertensi,
nigricans, occidentalis, orientalis, pachynema, perdita, pressula, subdermalis,
subnigricans, subterminalis, tentaculata, tenuissima, thompsoni, volcani.
Distribution. Eastern Panama and northern and central South America.
Fig. 2.5 contd
Fig. 2.5 A. Adult female Schistometopum thomense (Caeciliidae) and new-born from Ihla São Tomé,
Gulf of Guinea; the species is viviparous; photo by Ronald A. Nussbaum. B. Adult Chthonerpeton
indistinctum (Typhlonectidae) from southern South America; this species is terrestrial/semi-aquatic,
whereas other species of the family are fully aquatic; photo by John Measey. C-E. Adult
Scolecomorphus kirkii (Scolecomorphidae) from Tanzania; the eye, which rides on the base of the
tentacle, is shown under the skin and skull bones in the nearly resting position (C), in the nearly
maximally protruded position (D), and in the fully protruded position (E) in which the eye is carried
completely outside of the skull on the base of the tentacle; there is a pigmentless area of the skin over
the track of the eye as it moves back and forth with the tentacle, which presumably allows light to pass
through the skin and stimulate the retina; see O’Reilly et al. (1996) for details; photos by Daniel Boone.
Colour
Figure
Fig. 2.6 contd

Remarks. Caecilia is the largest genus of caecilians in the New World and
has a broad distribution. Over half the species were described by E. H.
Taylor, and most are poorly characterised and delimited. Surprisingly, there
have been no new species described and little taxonomic work on the group
since 1989 despite the clear need for the latter. A helpful treatment of the
Caecilia of Colombia is given by Lynch (1999). Summers and Wake (2001)
redescribed the holotype of C. volcani. C. isthmica was accidentally omitted
from our previous treatment (although counted in the total number of
species).
4. Dermophis Peters 1879
Type species. Siphonops mexicanus Duméril & Bibron 1841, by subsequent
designation of Noble (1924).
mesethmoid covered or exposed; no splenial teeth; secondary grooves
present; scales present; tentacular opening closer to eye than to external
naris; no unsegmented terminal shield; no narial plugs; no diastema
between vomerine and palatine teeth; no terminal keel.
Content. 7 species: costaricensis, glandulosus, gracilior, mexicanus, oaxacae,
occidentalis, parviceps.
Distribution. Southern Mexico south to northwestern Colombia.
Remarks. Following our 1989 comment that some of the species of
Dermophis considered invalid by Savage and Wake (1972) were valid,
Savage and Wake (2001) resurrected four species from the synonymies they
had previously proposed.
5. Gegeneophis Peters 1879
Type species: Epicrium carnosum Beddome 1870, by original monotypy.
mesethmoid not exposed dorsally; splenial teeth present; secondary
grooves present; scales present; tentacular opening midway between eye
and external naris; no unsegmented terminal shield; narial plugs on tongue;
no diastema between vomerine and palatine teeth; terminal keel present or
absent.
Content. 8 species: carnosus, danieli, fulleri, krishni, madhavai, nadkarnii,
ramaswamii, seshachari.
Fig. 2.6 contd
Fig. 2.6 A. Adult female Ichthyophis kohtaoensis (Ichthyophiidae) from Thailand guarding her clutch of
early-stage embryos; the species is oviparous with indirect development; the nest is terrestrial; the
hatchling larvae make their way to nearby streams where they grow and eventually metamorphose into
terrestrial subadults; photo by Alex Kupfer. B. Adult female Boulengerula boulengeri (Caeciliidae) from
Tanzania guarding her early-stage embryos in a terrestrial nest; the species is oviparous with direct
development (no larval stage); photo by Alex Kupfer. C. Adult Boulengerula taitanus (Caeciliidae) from
the Taita Hills, Kenya, guarding her early-stage embryos in a terrestrial nest; the species is oviparous
with direct development; photo by Alex Kupfer.
Distribution. India.
Remarks. Five new species of Gegeneophis have recently been described
from Maharashtra (Ravichandran et al. 2003; Giri et al. 2003) Karnataka
(Pillai and Ravichandran 1999; Bhatta and Srinivasa 2004) and Goa (Bhatta
and Prasanth 2004). Giri et al. (2003) also revised the generic diagnosis. No
uniquely derived traits are known for this genus.
6. Geotrypetes Peters 1880
Type species. Caecilia seraphini Duméril 1859, by original monotypy.
Diagnosis. Caeciliids with eye not covered with bone; temporal fossae
present; mesethmoid exposed dorsally; splenial teeth present; secondary
grooves present; scales present; tentacular opening closer to external naris
than to eye; no unsegmented terminal shield; narial plugs present on
tongue; no diastema between vomerine and palatine teeth; no terminal keel.
Content. 3 species: angeli, pseudoangeli, seraphini.
Distribution. Equatorial West Africa and Bioko Island.
Remarks. Nussbaum and Pfrender (1998) noted that Schistometopum
garzonheydti from Bioko is a junior synonym of Geotrypetes seraphini.
Geotrypetes seraphini have a distinctively shaped os basale (pers. obs.) and
unique arrangements of cranial muscles and anterior trunk muscles (Sheps
et al. 1997; Wilkinson, unpublished) which, if present in the other species,
would provide strong support for the monophyly of the genus.
7. Grandisonia Taylor 1968
Type species: Hypogeophis alternans Stejneger 1893, by original designation.
grooves present on more than half of the primary annuli, may be missing
on some anterior primaries; scales present; tentacular opening variable in
position, may be closer to external naris than to eye, nearly midway
between eye and external naris, or slightly closer to eye; no unsegmented
terminal shield; narial plugs present on tongue; no diastema between
vomerine and palatine teeth; no terminal keel.
Content. 4 species: alternans, brevis, larvata, sechellensis.
Distribution. Seychelles Archipelago.
Remarks. Grandisonia diminutiva is based on juvenile specimens of G.
sechellensis (Nussbaum, unpublished). The small “tail” that Taylor (1968)
considered to be diagnostic of G. diminutiva does not exist, and the rest of
the characteristics of the type series fall well within the range of G.
sechellensis.
8. Gymnopis Peters 1874
Type species: Gymnopis multiplicata Peters 1874, by original monotypy.
Diagnosis. Caeciliids with eye covered by bone; no temporal fossae;
grooves present; scales present; tentacular opening closer to eye than to
external naris; no unsegmented terminal shield; no narial plugs; no

diastema between vomerine and palatine teeth; no terminal keel.
Content. 2 species: multiplicata, syntrema.
Distribution. Guatemala south to Panama.
Remarks. The complicated history of the taxonomy of Gymnopis syntrema
was reviewed by Nussbaum (1988). Wake (in Savage and Wake 2001: 52)
indicated that she ‘disagrees with Nussbaum’s concept of Gymnopis as it
relates to G. syntremus [sic]’ and would treat the issue elsewhere.
9. Herpele Peters 1879
Type species: Caecilia squalostoma Stutchbury 1834, by original monotypy.
mesethmoid slightly visible or not dorsally; splenial teeth present;
secondary grooves present; scales present; tentacular opening closer to
external naris than to eye; no unsegmented terminal shield; narial plugs
present; no diastema between vomerine and palatine teeth; no terminal
keel.
Content. 2 species: multiplicata, squalostoma.
Distribution. Equatorial West Africa, including the Gulf of Guinea island,
Bioko (Fernando Po).
Remarks. The status of Herpele multiplicata, a species known only from a
holotype specimen that is now lost, was recently reviewed by Wilkinson et
al. (2003a) who wrongly gave the date of description of H. squalostoma as
1859 instead of 1834. H. squalostoma has an unusual arrangement of its
systemic arches that is unique among vertebrates (Wilkinson 1992b), which,
if present also in H. multiplicata, would provide good evidence of
monophyly of the genus.
10. Hypogeophis Peters 1879
Type species: Coecilia rostrata Cuvier 1829, by subsequent designation of
Parker (1958).
Diagnosis. Caeciliids with eye in socket, not under bone; no temporal
fossae; mesethmoid not exposed dorsally; secondary grooves present,
confined to posterior third or less of body; scales present; tentacular
opening far forward, closer to external naris than to eye; no unsegmented
terminal shield; narial plugs present; no diastema between vomerine and
palatine teeth; no terminal keel.
Content. l species: rostratus.
11. Idiocranium Parker 1936
Type species: Idiocranium russelli Parker 1936, by original designation and
monotypy.
Diagnosis. Caeciliids with eye not under bone; no temporal fossae;
mesethmoid widely exposed dorsally; frontals reduced, not in contact with
maxillaries; nasal in contact with squamosal; splenial teeth present;
secondary grooves present; scales present; tentacular opening closer to

external naris than to eye; no unsegmented terminal shield; narial plugs
present; no diastema between vomerine and palatine teeth; no terminal keel.
Content. l species: russelli.
Distribution. Cameroon.
12. Indotyphlus Taylor 1960
Type species: Indotyphlus battersbyi Taylor 1960, by original designation and
monotypy.
external naris; no unsegmented terminal shield; narial plugs on tongue; no
Content. 2 species: battersbyi, maharashtraensis.
Remarks. Giri et al. (2004) recently described the second known species of
Indotyphlus, noting that the narial plugs are not particularly small in either
species, a feature we previously included in the generic diagnosis. We know
of no uniquely derived traits.
Distribution. India, northern Western Ghats.
13. Luetkenotyphlus Taylor 1968.
Type species: Siphonops brasiliensis Lütken, 1852, by original designation
and monotypy.
Diagnosis. Caeciliids with eye not under bone; no temporal fossae; dorsal
exposure of mesethmoid unknown; no splenial teeth; no secondary
grooves; no scales; tentacular opening closer to eye than to external naris;
an unsegmented terminal shield; no narial plugs; premaxillary-maxillary
series of teeth short, not extending posterior to the choanae; no diastema
between vomerine and palatine teeth; a diastema between anterior ends of
the two series of vomerine teeth in adults; no terminal keel.
Content. l species: brasiliensis.
Distribution. Argentina, Brazil, Paraguay.
Remarks. Nussbaum (1986) and Nussbaum and Wilkinson (1989) used the
spelling “Lutkenotyphlus” for this genus, because we assumed that Lütken
is not a German word. This assumption was based on the fact that Christian
Frederik Lütken, for whom the genus is named, is Danish, and his ancestry
is also Danish. However, under a strict interpretation of the rules of
zoological nomenclature—when it doubt consider a word with an umlaut
to be a German word—our earlier spelling appears to have been an
unjustified emendation.
14. Microcaecilia Taylor 1968
Type species: Dermophis albiceps Boulenger 1882, by original designation.
mesethmoid not exposed dorsally; no splenial teeth; secondary grooves
usually present, absent in one species; scales present; tentacular opening

closer to eye than to external naris; no unsegmented terminal shield; no
narial plugs; no diastema between vomerine and palatine teeth; terminal
keel present or absent.
Content. 5 species: albiceps, rabei, supernumeraria, taylori, unicolor.
Distribution. Ecuador, French Guiana, Guyana, Surinam, Venezuela.
Remarks. Microcaecilia lacks any known uniquely derived features
supporting its monophyly. Our previous diagnosis included the absence of
a terminal keel, but this is present in at least M. unicolor (Wilkinson,
unpublished). Nussbaum (unpublished) has examined specimens from
northern Brazil that appear to be assignable to Microcaecilia.
15. Mimosiphonops Taylor 1968
Type species: Mimosiphonops vermiculatus Taylor 1968, by original
designation and monotypy.
Diagnosis. Caeciliids with eye in socket, not under bone; presence or
absence of temporal fossae and dorsal exposure of mesethmoid unknown,
probably as in Siphonops; splenial teeth present; no secondary grooves; no
scales; tentacular opening nearly equidistant between eye and external
naris; an unsegmented terminal shield; no narial plugs; a diastema between
Content. 2 species: reinhardti, vermiculatus.
Distribution. southern Brazil.
Remarks. Wilkinson and Nussbaum (1992) placed Pseudosiphonops in the
synonymy of Mimosiphonops and P. ptychodermis in the synonymy of M.
vermiculatus. Wake (2003) gave separate accounts for Pseudosiphonops and
Mimosiphonops in her review of caecilian osteology but did not comment on
the earlier proposed synonymy and presented no evidence that would
count against it. Mimosiphonops lacks known uniquely derived traits. Within
the siphonoforms (Mimosiphonops, Luetkenotyphlus, Siphonops) the relatively
anterior tentacle position, more strongly recessed mouths, and strong
diastema between the vomerine and palatine teeth are probably derived.
16. Oscaecilia Taylor 1968
Type species: Caecilia ochrocephala Cope 1866, by original designation.
mesethmoid exposed dorsally; splenial teeth present; secondary grooves
present; scales present; subdermal scales present or absent; tentacular
opening directly below external naris, closer to naris than to eye; no
unsegmented terminal shield; narial plugs present; no diastema between
vomerine and palatine teeth; no terminal keel; teeth relatively few and
large, replaced alternately in groups; vomeropalatine tooth row displaced
posteriorly, not parallel to premaxillary-maxillary tooth row, diverging from
the latter anteriorly forming an angle where the two rows meet rather than
a semicircle.
Content. 9 species: bassleri, elongata, equatorialis, hypereumeces, koepckeorum,

ochrocephala, osae, polyzona, zweifeli.
Distribution. Southern central and northern South America, Central
America (Costa Rica).
Remarks. Lahanas and Savage (1992) described O. osae from Costa Rica
since our last summary (Nussbaum and Wilkinson 1989).
17. Parvicaecilia Taylor 1968
Type species: Gymnopis nicefori Barbour 1924, by original designation.
Diagnosis. Caeciliids with eye not under bone; presence or absence of a
temporal fossae and dorsal exposure of mesethmoid unknown; no splenial
teeth; secondary grooves present; scales present; tentacular opening closer
to eye than to external naris; no unsegmented terminal shield; no narial
plugs; premaxillary-maxillary series of teeth short, not extending posterior
to the choanae; no diastema between the vomerine and palatine teeth; no
terminal keel.
Content. 2 species: nicefori, pricei.
Distribution. Colombia.
Remarks. We know of no uniquely derived traits of this poorly known
genus.
18. Praslinia Boulenger 1909
Type species: Praslinia cooperi Boulenger 1909, by monotypy.
mesethmoid not exposed dorsally; splenial teeth present; teeth small,
uniform in size, more than 50 per row, except for splenials; mouth terminal;
secondary grooves present; scales present; tentacular opening adjacent to
anterior edge of eye; no unsegmented terminal shield; no narial plugs; no
Content. l species: cooperi.
19. Schistometopum Parker 1941
Type species: Dermophis gregorii Boulenger 1894, by original designation.
fossae; mesethmoid exposed dorsally; splenial teeth present; secondary
external naris; no unsegmented terminal shield; no narial plugs; no
Content. 2 species: gregorii, thomense.
Distribution. Kenya, Tanzania, Gulf of Guinea islands.
Remarks. Nussbaum and Pfrender (1998) noted that S. ephele is a
geographic variant of S. thomense; S. brevirostre is a junior synonym of S.
thomense; and S. garzonheydti is a junior synonym of Geotrypetes seraphini.
Gower and Wilkinson (2002) suggested that the species of Schistometopum
share a uniquely derived phallus ornamentation, and there is strong

support for monophyly from molecular data (Wilkinson et al., 2003b).
20. Siphonops Wagler 1828
Type species: Caecilia annulata Mikan 1820, by original monotypy.
fossae; mesethmoid exposed dorsally; no splenial teeth; no secondary
grooves; no scales; tentacular opening closer to eye than to external naris;
an unsegmented terminal shield; no narial plugs; no diastema between
Content. 5 species: annulatus, hardyi, insulanus, leucoderus, paulensis.
Distribution. Argentina, Bolivia, Brazil, Colombia, Ecuador, Guyana,
Paraguay, Peru, Venezuela, and probably Uruguay, Surinam, and French
Guiana.
Remarks. We know of no uniquely derived traits. The three species
annulatus, leucoderus, paulensis have a distinctive and presumably derived
colour pattern with a blue background and whitish annular ring, but this is
shared with the species of Mimosiphonops suggesting that Siphonops may be
paraphyletic.
21. Sylvacaecilia Wake 1987
Type species: Geotrypetes grandisonae Taylor 1970, by original designation
and monotypy.
external naris; no unsegmented terminal shield; narial plugs present; no
Content. l species: grandisonae.
Distribution. Ethiopia.
F. Family Typhlonectidae Taylor 1968 (Fig. 2.5 B)
Diagnosis. Gymnophiona with the same number and arrangement of skull
bones as caeciliids, but pseudoectopterygoid never present and
mesethmoid always covered by frontals; temporal fossae present; eye never
under bone; splenial teeth present; teeth monocuspid; m. interhyoideus
posterior short; tentacle small; choanae large with well developed valves;
narial plugs present; relatively large cloacal disk; viviparous; embryonic
gills fused into a large, sac-like structure on each side; undivided primary
annuli only, or some primary annuli with pseudosecondary grooves; aortic
arches proximal to the heart fused into an elongate truncus arteriosus; atrium
partially divided externally.
Content. 5 genera and 12 species.
Distribution. South America.
1. Atretochoana Nussbaum and Wilkinson 1995.
Type species: Typhlonectes eiselti Taylor 1968, by original designation and

monotypy.
Diagnosis. Typhlonectids with sealed choanae; no lungs; no pulmonary
blood vessels; postcranial jaw articulation; posteriorly directed and
elongate stapes; novel stapedial muscle; the tentacular aperture
intermediate in position between eye and external naris; tentacular groove
not covered with bone; body laterally compressed with a middorsal fin;
unknown habit, but suspected to be lotic-torrential.
Content. 1 species: eiselti.
Distribution. ‘South America’.
Remarks. The holotype of A. eiselti lacks detailed locality data, but is
labelled “South America”. Wilkinson et al. (1998) reported the second
known specimen of A. eiselti, which is also without data but suspected to
be from Brazil.
2. Chthonerpeton Peters 1879
Type species: Siphonops indistinctus Reinhardt and Lütken 1861, by
monotypy.
Diagnosis. Typhlonectids with the tentacular aperture intermediate in
position between eye and external naris; tentacular groove not covered with
bone; foetal gills attaching laterally to the nuchal region, the two gill bases
well separated dorsally; no lateral compression of the body; no middorsal
ridge or free fold (fin); left lung rudimentary; external naris ovate; choanal
valve aperture along entire length of valve; cloacal disk subcircular;
semiaquatic habit.
Content. 8 species: arii, braestrupi, exile, indistinctum, noctinectes, onorei,
perissodus, viviparum.
Distribution. Northern Argentina, Brazil, Ecuador, Uruguay.
Remarks. The two species C. arii (Cascon and Lima-Verde 1994) and C.
noctinectes (Silva et al. 2003) were described since 1989. Based on specimens
in Museum collections, there appears to be a number of undescribed
Chthonerpeton (Wilkinson, unpublished). Uniquely derived traits of
Chthonerpeton are unknown, but the type species, C. indistinctum lacks a
distinct m. rectus lateralis, a highly unusual condition unknown in other
caecilians. If true of the other species this would provide strong support for
the monophyly of the genus.
3. Nectocaecilia Taylor 1968
Type species: Chthonerpeton petersii Boulenger 1882, by original designation.
Diagnosis. Typhlonectids with the tentacular aperture close behind external
naris; tentacular groove partially roofed by bone in adults; fetal gills
attaching dorsolaterally, the two gill bases slightly separated middorsally;
no lateral compression of the body; no dorsal free fold or ridge; left lung
well developed; subcircular cloacal disk; external naris subtriangular;
choanal valve aperture along entire length of the valve; semiaquatic habit.
Content. 1 species: petersii.

Distribution. Venezuela.
Remarks. Wilkinson (1996b) placed N. haydee (Roze 1963) in the synonymy
of Typhlonectes natans.
4. Potomotyphlus Taylor 1968
Type species: Caecilia kaupii Berthold 1859, by original designation.
Diagnosis. Typhlonectids with the tentacular aperture close behind external
naris; tentacular groove partially roofed with bone in adults; foetal gills
unknown; body laterally compressed; middorsal free fold or ridge present;
left lung well developed, dilated, much wider than right lung; head small
relative to body; cloacal disk subcircular posteriorly with a narrower
anterior portion; external naris subtriangular; choana extremely large;
choanal valve with aperture restricted to a small funnel-like flap; aquatic
habit.
Content. l species: kaupii.
Distribution. Brazil, Ecuador, French Guiana, Peru, Venezuela.
5. Typhlonectes Peters 1879
Type species: Caecilia compressicauda Duméril and Bibron 1841, by
subsequent designation of Dunn (1942).
Diagnosis. Typhlonectids with tentacular aperture close behind external
naris; tentacular groove partially roofed by bone in adults; foetal gills
attaching dorsally, the two gill bases fused with no separation; body
laterally compressed, at least posteriorly, in adults; a middorsal ridge or
free fold present; left lung well developed; cloacal disk, subcircular;
external naris subtriangular; choanal valve aperture along full length of
valve; habit aquatic.
Content. 2 species: compressicauda, natans.
Distribution. Colombia, Peru, Venezuela, French Guiana, Guyana,
Amazonian Brazil.
Remarks. T. cunhai was described by Cascon et al. (1991) but its validity has
been questioned (Wilkinson and Nussbaum 1997). Based on recent
examination (Wilkinson, unpublished) we consider the holotype to be
indistinguishable from T. compressicauda and thus place T. cunhai in the
synonymy of T. compressicauda (Dumeril and Bibron, 1841). Wilkinson (1991;
1996c) provided formal synonymies for several species recognised by
Taylor (1968) but not included in our previous treatment (Nussbaum and
Wilkinson 1989). Wilkinson and Nussbaum (1999) identified nine uniquely
derived characters supporting monophyly of the genus.
2.4 ACKNOWLEDGEMENTS
We cannot individually acknowledge all of our many colleagues: field
workers, bench workers, researchers, curators, students and technicians
who have facilitated our work in one way or another, but we wish to
express our sincere thanks for these contributions without which most our
work would not be possible. We thank David Gower, Hendrik Müller and
Samantha Mohun for reviewing the manuscript and Alex Kupfer, Daniel
Boone, Peter Stafford, and John Measey for providing photographs. MW
gratefully acknowledges funding from University of Glasgow New
Initiatives Fund, the Museum and Zoology Research Funds of the Natural
History Museum, London, the Percy Sladen Memorial Trust and the NERC
(GST/02/832 and GR9/02881). RAN received grants from the National
Geographic Society and the U.S. National Science Foundation in support of
his field research with caecilians.
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Reproductive Biology and Phylogeny of Gymnophiona

Science Publishers, Enfield, NH, USA. 2006.

uncertainties that are currently a major obstacle to progress in caecilian

2.2 CAECILIAN PHYLOGENY

monophyletic group on the basis of cytogenetic data (Nussbaum and Ducey

separation of the rhinatrematids from all other caecilians was well

2.2.2 Non-rhinatrematids — Neocaecilia

2.2.5 Ichthyophiidae + Uraeotyphlidae — Diatriata

2.2.6 The Higher Caecilians — Teresomata

informally termed the advanced (Nussbaum 1991) or higher (e.g., San

posterior (Fig. 2.1), supports the monophyly of the Caeciliidae. However,

relationships that are well supported by molecular data, including the

species lists and synonymies. Where possible, we have indicated derived

ORDER GYMNOPHIONA RAFINESQUE 1814

Fig. 2.3 contd

Diagnosis. Rhinatrematids with two ceratobranchial arches in adults; more

Content. 5 species: interruptus, malabaricus, menoni, narayani, oxyurus.

Fig. 2.4 contd

Fig. 2.5 contd

2. Brasilotyphylus Taylor 1968

Fig. 2.6 contd

Fig. 2.6 contd

external naris; no unsegmented terminal shield; no narial plugs; no

secondary grooves present; scales present; tentacular opening closer to

usually present, absent in one species; scales present; tentacular opening

Content. 9 species: bassleri, elongata, equatorialis, hypereumeces, koepckeorum,

share a uniquely derived phallus ornamentation, and there is strong

Type species: Typhlonectes eiselti Taylor 1968, by original designation and

Content. 1 species: petersii.

2.5 LITERATURE CITED

Cascon, P. and Lima-Verde J. S. 1994. Uma nova especie de Chthonerpeton do nordeste

Guérin-Méneville, M. F. E. 1829-1844. Iconographie du Règne Animal de G. Cuvier

Nussbaum, R. A. 1981. Scolecomorphus lamottei, a new caecilian from West Africa

Wilkinson, M. 1996b. Resolution of the taxonomic status of Nectocaecilia haydee (Roze)

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