March 1989] HERPETOLOGICA 23

viridiflavus (Dumeril et Bibron, 1841) (Anura Hy- SELANDER, R. K., M. H. SMITH, S. Y. YANG, W. E. peroliidae) en Afrique centrale. Monit. Zool. Itali- JOHNSON, AND J. B. GENTRY. 1971. Biochemical ano, N.S., Supple. 1:1-93. polymorphism and systematics in the genus Pero- LYNCH, J. D. 1966. Multiple morphotypy and par- myscus. I. Variation in the old-field mouse (Pero- allel polymorphism in some neotropical frogs. Syst. myscus polionotus). Stud. Genetics IV, Univ. Texas Zool. 15:18-23. Publ. 7103:49-90. MAYR, E. 1963. Species and Evolution. Har- SICILIANO, M. J., AND C. R. SHAW. 1976. Separation vard University Press, Cambridge. and localization of enzymes on gels. Pp. 184-209. NEI, M. 1972. Genetic distance between popula- In I. Smith (Ed.), Chromatographic and Electro- tions. Am. Nat. 106:283-292. phoretic Techniques, Vol. 2, 4th ed. Williams Hei- RESNICK, L. E., AND D. L. JAMESON. 1963. Color nemann' Medical Books, London. polymorphism in Pacific treefrogs. Science 142: ZIMMERMAN, H., AND E. ZIMMERMAN. 1987. Min- 1081-1083. destanforderungen fur eine artgerechte Haltung SAVAGE, J. M., AND S. B. EMERSON. 1970. Central einiger tropischer Anurenarten. Zeit. Kolner Zoo American frogs allied to Eleutherodactylusbrans- 30:61-71. fordii (Cope): A problem of polymorphism. Copeia 1970:623-644. Accepted: 12 March 1988 Associate Editor: John Iverson SCHIOTZ, A. 1971. The superspecies Hyperoliusvir- idiflavus (Anura). Vidensk. Medd. Dansk Natur- hist. Foren. 134:21-76.

Herpetologica,45(1), 1989, 23-36 ? 1989 by The Herpetologists'League, Inc.

ON THE STATUS OF NECTOCAECILIA FASCIATA TAYLOR, WITH A DISCUSSION OF THE PHYLOGENY OF THE (AMPHIBIA: )

MARK WILKINSON Museum of Zoology and Department of Biology, University of Michigan, Ann Arbor, MI 48109, USA

ABSTRACT: Nectocaecilia fasciata Taylor is a junior synonym of Chthonerpetonindistinctum Reinhardt and Liitken. The range of Nectocaecilia is restricted to northern South America. A hypothesis of phylogenetic relationships among the four genera of the Typhlonectidae is proposed, and alternative hypotheses are evaluated. The recently established genus Pseudotyphlonectes and other higher taxonomic categories proposed by Lescure et al. (1986) for the Typhlonectidae are invalid. Pseudotyphlonectes is a junior synonym of Typhlonectes Peters. It is suggested that phy- logenetic studies of the Gymnophiona should proceed independently of taxonomic considerations until a compelling hypothesis is accepted. Key words: Nectocaecilia; Chthonerpeton; Potomotyphlus; Typhlonectes; Pseudotyphlo- nectes; Phylogeny; ; Typhlonectidae; Gymnophiona

RECENTLY, within a single year, three sults are essentially in agreement with his, separate taxonomic treatments and hy- resulting in a single taxonomic change. potheses of phylogeny of the Gymno- Laurent (1986) and Lescure et al. (1986) phiona were published. These three works both produced hypotheses of intra-generic differed in scope, methodology, and re- relationships within the Gymnophiona. sults. Duellman and Trueb (1986) used cla- Laurent's methodology was not made clear, distic techniques to analyze relationships but that of Lescure et al. was claimed to among families and subfamilies. They re- be cladistic. The latter authors' phyloge- lied heavily on characters that Nussbaum netic hypothesis was coupled with a pro- (1977, 1979) had reported, and their re- posal for major taxonomic rearrangements 24 HERPETOLOGICA [Vol. 45, No. 1 within the Gymnophiona. The phyloge- positioned perpendicular to the body sur- nies and taxonomic treatments of these au- face at the level of the third nuchal groove thors are markedly different from each or were inserted into the body at this level. other and from that of Duellman and Counts made in these ways did not differ Trueb (1986). Nussbaum and Wilkinson from counts made from radiographsof the (in press) critically evaluated the three same specimens without markersusing the phylogenetic hypotheses and concluded visible constriction corresponding to the that the proposed phylogenies of Laurent third nuchal groove to delimit the nuchal (1986) and Lescure et al. (1986) are not region. These counts are frequently not supported by the evidence and that too whole numbers, presumably because the little is presently known about variation in external muscular sheath, to which the characteristics to justify taxo- dermis attaches, is itself not strongly at- nomic rearrangements at or above the fa- tached to deeper vertebral structures milial level. (Gaymer, 1971; Naylor and Nussbaum, The taxonomy of the Gymnophiona is 1980), and relative movement between the also unstable at the generic and specific vertebral column and external annular or levels. Research is constantly revealing taxa nuchal grooves is possible. Most frequent- that are invalid and errors in established ly, fractional numbers of nuchal vertebrae taxonomic treatments (Nussbaum, 1985, are encountered in preserved specimens 1986a, in press; Nussbaum and Gans, 1981; that have the nuchal region bent or oth- Savage and Wake, 1972; Wilkinson, 1988). erwise contorted. At present, genera are poorly defined, and Postcloacal vertebral counts were made for many there can be little guarantee that from radiographs using pins inserted into they are natural. Whilst such problems the cloacal aperture perpendicular to the persist, hypotheses of intra-generic rela- body surface. The numbers of postcloacal tionships are flawed by their reliance on vertebrae may be subject to artifactual possibly artificial units. Thus, it is impor- variation, increasing with dehydration as tant to redefine natural units that may then the soft body parts shrink around the ver- serve as a basis for meaningful comparison. tebral column (Nussbaum, in press). The In this paper, I point out one such prob- range of postcloacal vertebrae in Chtho- lem in the genus Nectocaecilia. By re- nerpeton is increased by such variation, stricting the concept of Nectocaecilia, I but for the purposes of this paper, it was evaluate its position within the Typhlo- not necessary to discriminate between nectidae and present evidence for an al- counts derived from well preserved or de- ternative phylogenetic hypothesis for the hydrated specimens. typhlonectid genera. I argue that this hy- Measurementswere made to the nearest pothesis should be preferred over those 0.1 mm with dial calipers, except total proposed by Laurent (1986) and by Les- lengths, measured to the nearest 1 mm by cure et al. (1986). I cannot evaluate the stretching specimens along a meter rule. work of Duellman and Trueb (1986) with In order to evaluate the results and the evidence presented here, because they methodology of Lescure et al. (1986), their did not address the interrelationships of data for the Typhlonectidae were reana- the typhlonectid genera. lyzed with a computerized parsimony al- gorithm (PAUP, version 2.4: Swofford, 1985). All charactersthat varied acrossthe METHODS typhlonectid taxa considered by Lescure Counts of vertebrae in the nuchal region et al. were included in the reanalysis.These were made from radiographs. Specimens characters were recoded in binary form in which the nuchal region was well de- using the morphoclines of Lescure et al., fined and clearly delimited by the third except for their annulation character for nuchal groove (= first primary groove) which polarization of the character states were used. For some specimens, pins were was not possible. For this character, the used as radio-opaque markers and were states reported by Lescure et al. in the March 1989] HERPETOLOGICA 25

"remarques sur segments succesion an- only the number of primary annuli and tero-posterieure" section of their data ma- tooth counts reportedby Taylor are similar trix were assigned symbols and run unor- to my data for IMZT An-786 (Table 1). dered. It would seem that Taylor's type de- The topology of the cladogram of Les- scription was based on a different speci- cure et al. (1986) was specified and PAUP men, and if this was the case, Nectocaecilia was used to distribute the character states fasciata would be a nomen dubium until upon this cladogram so as to minimize ho- either the unknown "holotype" or new moplasy. The ALLTREES routine of specimens corresponding to Taylor's de- PAUP was used to generate all possible scription are discovered. Taylor's data for cladogram topologies and to identify the the "holotype" of N. fasciata includes a ones that best fit the data. PAUP provided 1.0 mm projection of the snout. This is consistency indices for each of these trees anomalous because the Typhlonectidae is and for the cladogram of Lescure et al. a family typified by strongly recessed which facilitated comparison. mouths, and a typhlonectid as large as the holotype (390 mm total length) should have STATUSOF NECTOCAECILIAFASCIATA a snout projection greater than 3.0 mm. If Taylor (1968) established the genus Taylor actually examined a specimen that Nectocaecilia to receive Chthonerpeton corresponds to his description of N. fas- petersii Boulenger, a species then known ciata, then it is surprising that this speci- only from the holotype and for which the men was referred by Taylor to the Typh- only available locality was the Upper Am- lonectidae, and that the minimally recessed azon; C. haydee Roze, from Venezuela; mouth, which would be its most distinctive and two new forms, N. ladigesi, from Rio feature, was neither commented upon nor Moju, Brazil, and N. fasciata from Buenos used as a diagnostic character by Taylor. Aires, Argentina. These taxa were linked In the Museo Civico di Storia Naturale, by the unique combination of characters: Genova, there is a specimen of Typhlo- sub-triangular nares, tentacular apertures nectes natans (MSNG 379/A) which was far forward behind the nares (as in Typh- received as a gift from the University of lonectes and Potomotyphlus), and the ab- Torino and which agrees fairly well with sence or vague trace of a fin in adults (well Taylor's description of Nectocaecilia fas- developed in Typhlonectes and Potomo- ciata excepting the projection of the snout typhlus, absent in Chthonerpeton). Taylor and the tooth and annular counts (Table (1970) described a further species of Nec- 1). This specimen is a male and the dorsal tocaecilia (N. cooperi), from Rio Magda- "fin" is basically restricted to a ridge, de- lena, Colombia, which Wilkinson (1988) veloping into a free fold only above the showed was based on a single aberrant terminus. specimen of Typhlonectes natans (Fisch- The IMZT collections contain, in addi- er). tion to the single specimen of Chthoner- Taylor (1968:269) distinguished Nec- peton indistinctum, two more specimens tocaecilia fasciata from the other species of Typhlonectes natans and five of Si- of Nectocaecilia by its low number of pri- phonops annulatus (Mikan). In the latter mary annuli, and he reported that the ho- species, the projection of the snout is far lotype and only known specimen is in the less than in any typhlonectid and is com- Museo di Zoologia, University of Torino, parable to that reported by Taylor for Nec- IMZT 2817 (since recatalogued as IMZT tocaecilia fasciata. I suspect that Taylor An-786; see Elter, 1981). I recently ex- examined all of these specimens, including amined the small collection of the T. natans now in Genova, and con- in Torino and found that IMZT An-786 is fused his data such that his description of a specimen of Chthonerpeton indistinc- N. fasciata was a composite of data re- tum Reinhardt and Liitken (Figs. IA, 2A). corded from C. indistinctum with its low This specimen differs significantly from number of annuli, T. natans with its large Taylor's type description of N. fasciata; sub-triangular nares and closely positioned 26 HERPETOLOGICA [Vol. 45, No. 1

A B

cQDW

FIG. 1.-Whole body photographs of representative typhlonectids. (A) Chthonerpeton indistinctum, IMZT An-786, Total length (TL) = 175 mm. (B) Nectocaecilia petersii, AMNH A100589, TL = 505 mm. (C) Potomotyphius kaupii, MCZ 96758, TL = 605 mm. (D) Typhlonectes natans, CAS 153871, TL = 570 mm. (E) Typhlonectes compressicaudus, CM 90091, TL = 464 mm. Note the development of middorsal free folds, modification of the terminal shield and lateral compression of the body in (C), (D), and (E) and their absence in (A) and (B).

tentacular apertures, and S. annulatus with is IMZT An-786, and so I consider Nec- its feebly projecting snout. These features tocaecilia fasciata Taylor to be a junior distinguished Taylor's N. fasciata from all synonym of Chthonerpeton indistinctum other caecilians. Reinhardt and Liitken. The other nominal If this interpretation is accepted, Nec- species of Nectocaecilia are known only tocaecilia fasciata cannot be a nomen du- from South America no farther south than bium. The named holotype of N. fasciata the Amazon drainage, and the anomaly of March 1989] HERPETOLOGICA 27 e e

n n t ~~~~~~~~~~~~t A B

n t e ~~~~

#017,~ ~ ~

n tn tt E

FIG. 2.-Left side of the head of representative typhlonectids showing relative positions of the eye (e), tentacular aperture (t), and naris (n). (A) Chthonerpeton indistinctum, IMZT An-786, Head length (HL = distance from snout tip to first nuchal groove) = 8.6 mm. (B) Nectocaecilia petersii, AMNH A100589, HL = 13.3 mm. (C) Potomotyphlus kaupii, MCZ 96758, HL = 12.4 mm. (D) Typhlonectes natans, CAS 153871, HL = 23.3 mm. (E) Typhlonectes compressicaudus, CM 90090, HL = 17.5 mm. Note the strong projection of the snout beyond the recessed mouth that is typical of all typhlonectids, and the naris shape; ovate in (A), sub-triangular in (B), (C), (D), and (E). a congener as far south as Argentina is now close behind them. By restricting consid- removed. eration of Nectocaecilia to N. petersii only, Of the remaining species of Nectocae- it becomes possible to evaluate the phy- cilia, the validity of N. haydee is uncertain logenetic position of Nectocaecilia within and N. ladigesi is a Typhlonectes (Wil- the Typhlonectidae. kinson, unpublished), but N. petersii, the type species, is a distinctive, undoubtedly PHYLOGENYOF THE TYPHLONECTIDAE valid species which has no middorsal ridge The family Typhlonectidae consists of nor free fold and has sub-triangular narial four genera. Nectocaecilia seems inter- apertures with the tentacular apertures mediate in its most apparent characteris- 28 HERPETOLOGICA [Vol. 45, No. 1

TABLE 1.-Morphometric and meristic data reported Chthonerpeton the sister taxon of this as- for Nectocaecilia fasciata by Taylor (1968:269-271) semblage, and the second composed of Po- and original data for IMZT An-786 (Chthonerpeton indistinctum) and for MSNG 379/A (Typhlonectes tomotyphlus and Typhlonectes. natans) believed to have been confused in Taylor's description of N. fasciata. All measurements in mm. MONOPHYLY OF NECTOCAECILIA, N. fasciata C. indis- T. POTOMOTYPHLUS,AND TYPHLONECTES (data tinctum natans from (IMZT (MSNG Neither of the positions of the tentacular Taylor) An-786) 379/A) aperture found in the Typhlonectidae cor- Total length 390 175 385 responds exactly to the positions found in Number of annuli 76 74 89 any other caecilian genus, and thus both Width 22 6.5 16* Length divided by width 17.7 26.9 24.1 positions might be derived relative to other Head width 14 5 12 Gymnophiona. The intermediate position Head length to 1st nuchal in Chthonerpeton is more closely ap- groove 17 8.6 16.3 proached in other forms. Amongst neo- Head length to 2nd nuchal tropical genera, a position of the groove 23 11.6 21.3 tentacular Head length to 3rd nuchal aperture intermediate between the eye and groove 28.2 13.6 28.2 naris is typical (exceptions to this are Rhi- Projection of snout natrema, , Caecilia, and Os- beyond mouth 1.0 1.8 3.2 caecilia), but it is usually closer to the lip Tentacular aperture to naris 0.7 0.12 0.7 (e.g., Dermophis) or closer to the eye and Premaxillary-maxillary more distant from the lip (e.g., Siphonops) teeth 28 27 than in Chthonerpeton. Thus the evidence Vomeropalatine teeth 27 24 suggests that the position of the tentacular Dentary teeth 24 26 aperture in Splenial teeth 6 6 Chthonerpeton may be prim- itive for the Typhlonectidae. If both states * Width measurement affected by maceration of body musculature. are independently derived from a differ- ent state in the last common ancestor of tics between Chthonerpeton on the one these genera, then this would indicate hand, and Potomotyphlus and Typhlo- monophyly of the genus Chthonerpeton, nectes on the other. The lack of a mid- and of a group composed of the three gen- dorsal ridge or free fold in Chthonerpeton era Nectocaecilia, Potomotyphlus, and and Nectocaecilia (Fig. 1A,B) is probably Typhlonectes. If the condition in Chtho- primitive. The presence of well developed nerpeton is primitive relative to the con- free folds in females of Typhlonectes dition of the other typhlonectid genera, (ridges or free folds in males) and in both this would still constitute evidence of sexes of Potomotyphlus (Fig. 1C-E) is monophyly of the Nectocaecilia-Poto- probably a shared derived feature (found motyphlus-Typhlonectes group, but not in the adults of no other caecilian genus). of monophyly of Chthonerpeton. The pos- This is evidence that Potomotyphlus and sibility that the intermediate position of Typhlonectes are sistertaxa. Nectocaecilia the tentacular aperture is derived from the shares with Potomotyphlus and Typhlo- more anterior position, which would con- nectes (Fig. 2B-E) tentacular apertures stitute evidence only for monophyly of that are close behind the nares and sub- Chthonerpeton, seems unlikely. Thus, I triangular narial apertures. In Chthoner- interpret the position of the tentacular ap- peton (Fig. 2A), the tentacular apertures erture in Nectocaecilia, Potomotyphlus, are more nearly equidistant between the and Typhlonectes as a shared derived eyes and nares, and the narial apertures character indicating monophyly of this are more ovate and elongate. group. Nussbaum (1977, 1979) used an ar- All of the evidence that I discuss indi- gument from ontogeny to polarize these cates monophyly of two groups: the first character states. Duellman and Trueb including the three genera Nectocaecilia, (1986) simplified Nussbaum's (1979) treat- Potomotyphlus, and Typhlonectes, with ment of this character. Both of these in- March 1989] HERPETOLOGICA 29

terpretations are consistent with that pro- indicating monophyly of this family. posed here. Within the family, Chthonerpeton has the The same interpretation can be applied smallest choanal aperturesand the smallest to the shape of the narial apertures. Both and deepest choanal valves. Larger choa- conditions found in the Typhlonectidae nae and larger valves are present in the might be independently derived from some other three genera, forming a graded series different form, or any one state could be of increasing size. I interpret this as two primitive relative to the other. Among oth- parallel evolutionary transformationseries er caecilians, narial aperture shape is typ- in which Chthonerpeton has the most ically subcircular, but in some species of primitive states (the smallest choanae and Caecilia and Siphonops, they may be smallest valves), Potomotyphlus and slightly ovate and elongated, approaching Typhlonectes share the most derived states the condition in Chthonerpeton, which (extremely enlarged choanae and valves), may thus be primitive for the Typhlonec- and Nectocaecilia has intermediate states tidae. That ovate narial apertures might (moderatelyenlarged choanae and valves). be derived from sub-triangularones seems Shared possession of the derived states of unlikely, and I conclude that sub-trian- these two characters (moderately or ex- gular narial aperturesare a shared derived tremely enlarged choanal apertures and trait indicating monophyly of the Necto- choanal valves) is evidence of monophyly caecilia -Potomotyphlus - Typhlonectes of the Nectocaecilia -Potomotyphlus- group. Typhlonectes group. These three genera In caecilians, the right lung is typically also share superficial placement of their small and the left is smaller still, often little choanal valves, and this is also a shared more than a small diverticulum. Within derived feature indicating their monophy- the Typhlonectidae, the lungs demonstrate ly. In Nussbaum's(1979) cladistic analysis, different degrees of elongation. Chtho- enlarged choanae were considered de- nerpeton clearly has the most primitive rived. His treatment, however, included pattern with a very short left lung and a only a single typhlonectid genus. I have right lung that stretchesposteriorly to about extended this treatmentby subdividingthis the level of the posterior end of the liver. character to reflect the different character Nectocaecilia, Potomotyphlus, and Typh- states present in the Typhlonectidae. lonectes share the derived features of a Without justification, Lescure et al. right lung that extends caudally much fur- (1986) considered four vertebrae in the ther than the liver, and a left lung that is nuchal region primitive and three or five also well developed. Both of these features derived. They reported that all of the are good evidence for the monophyly of typhlonectidtaxa that they consideredhave this group of three genera. three nuchal vertebrae; however, I find In non-typhlonectid caecilians, it is not (based on larger samples) that the number always easy to determine the presence of of nuchal vertebrae is variable within the choanal valves, because they may be con- Typhlonectidae (summarized in Table 2). cealed deep within the small choanae, Chthonerpeton has three nuchal verte- hence knowledge of their occurrence is brae, whereas Nectocaecilia, Potomotyph- very limited. I have observed them within lus, and Typhlonectes all have four or the neotropical genera Caecilia and Der- more. Also contraryto Lescureet al. (1986), mophis where they are small and deep, the dominant condition found in possible and Taylor (1968) reported them in Gran- neotropical outgroups for the Typhlonec- disonia from the Seychelles archipelago. tidae (Dermophis, Gymnopis, Siphonops, Choanal valves may well be widespread Caecilia, Epicrionops) is three nuchal ver- in the Gymnophiona but not visible in su- tebrae (Wilkinson, unpublished). perficial examination. In the Typhlonec- Lescure et al. (1986) reported the pres- tidae, the choanal aperturesare larger and ence of "tails", which they equated with the valves are larger and more superficial, postcloacalvertebrae, in Typhlonectes and and these are probably derived features their absence in Potomotyphlus and 30 HERPETOLOGICA [Vol. 45, No. 1

TABLE 2.-Summary of numbers of nuchal and posteloacalvertebrae in genera and species of the Typhlo- nectidae.

Numbers of vertebrae Nuchal Postcloacal Range x n Range x n

Chthonerpeton 2.8-3.8 3.24 40 0-3 1.71 33 Nectocaecilia 4.0-4.5 4.09 10 3.5-5 4.07 7 Potomotyphlus 4.0-5.0 4.06 15 4-5 4.2 14 Typhlonectes natans 4.0 4.0 35 3.5-6 4.31 38 T. compressicaudus 4.0-4.5 4.03 15 4-7 5.5 8

Chthonerpeton. There are good reasonsto of Nectocaecilia intermediate between prefer Nussbaum's(1979) equating of "true these two. tails" with postcloacal external segmen- tation supported by postcloacal vertebrae, because these "true tails" are probably not MONOPHYLY OF POTOMOTYPHLUS AND homologouswith postcloacalvertebrae that TYPHLONECTES occur in the absence of postcloacalexternal In addition to sharing middorsal ridges segmentation. My investigationsshow that or free folds, these two genera are sister within the Typhlonectidae only Chtho- taxa, as indicated by the size of their nerpeton may lack postcloacal vertebrae choanal apertures and choanal valves, (Table 2). This condition is also typical of modification of the body terminus, fea- neotropical caeciliids (an outgroup) indi- tures of the cloacal disc, lateral compres- cating that it is primitive for the Typhlo- sion of the body, and fully aquatic habits. nectidae. I interpret four nuchal vertebrae It is commonly stated that the typhlo- and several (>3) postcloacal vertebrae, in nectids are aquatic; however, only Typh- the absence of postcloacal external seg- lonectes and Potomotyphlus are truly mentation, as derived character states aquatic. Both Chthonerpeton and Nec- which are shared by, and are evidence of tocaecilia are more accurately described monophyly of, the Nectocaecilia-Poto- as semiaquatic (Nussbaum, 1986b; Tan- motyphlus-Typhlonectes group. ner, 1971). Correlated with the aquatic The gills of fetal typhlonectid caecil- habits of Typhlonectes and Potomotyph- ians attach to the nuchal region in gener- lus is lateral compression of at least the ically specific patterns that are readily posterior part of the body. In contrast, interpretable phylogenetically (R. A. Chthonerpeton, Nectocaecilia, and the Nussbaum, personal communication). In adults of all other caecilians have roughly Chthonerpeton, each gill attaches later- cylindrical or often slightly dorsoventrally ally, and the bases are widely separated compressed bodies. Nussbaum (1979) con- dorsally. This pattern of attachment is typ- sidered lateral compressionto be a derived ical of non-typhlonectid caecilians and is feature reflecting the aquatic specializa- probably primitive for the Typhlonecti- tion of the genus Typhlonectes. Duellman dae. Fetal Nectocaecilia have gills that at- and Trueb (1986) followed Nussbaum but tach dorsolaterallyand, as a consequence, erroneously considered lateral body the bases of the two gills are only slightly compression to be a characteristic of the separated middorsally. In the fetuses of Typhlonectidaeas a whole. I interpretboth both Typhlonectes compressicaudus (Du- the lateral compression and fully aquatic meril and Bibron) and T. natans, there is habits of Typhlonectes and Potomotyph- dorsal attachment of both gills, which are lus as shared derived characters. fused at their bases with no middorsalsep- Another common assertion is that the aration. The condition in Chthonerpeton body terminus surroundingthe cloacal ap- is considered the most primitive, that of erture forms a sucking or even prehensile Typhlonectes the most derived, and that organ used by the males for attachment to March 1989] HERPETOLOGICA 31 females during copulation (e.g., Taylor, most derived states, and both are shared 1968). The only reports of copulation in by Potomotyphlus and Typhlonectes. typhlonectids (Barrio, 1969; Murphy et al., If the gills of fetal Potomotyphlus (pres- 1977) lend no support to this speculation, ently unknown) are fused at their bases and there are good reasons, based on the rather than separate, this would constitute morphology of this region, to discount this additional evidence for monophyly of Po- notion (Wilkinson, unpublished). In many tomotyphlus and Typhlonectes. If, how- caecilians, the cloacal aperture lies within ever, the gills of fetal Potomotyphlus prove a bluntly rounded, externally unsegment- to be like those of Chthonerpeton, this ed, terminal portion of the body termed would be evidence contradicting my in- an unsegmented terminal shield. The clo- terpretation of monophyly of these two acal disc (the typically unpigmented re- genera. In either case, both T. compres- gion about the cloacal aperture that bears sicaudus and T. natans share the derived architecture continuous with the internal condition of fused gill bases, and this is architecture of the cloacal walls) lies with- compelling evidence that these two species in the shield. In non-typhlonectid caecili- share a more recent common ancestorthan ans, the disc is small and not or only scarce- either does with Nectocaecilia or with ly recessed. Within the Typhlonectidae, Chthonerpeton. this condition is most closely approached in Chthonerpeton and Nectocaecilia. In ALTERNATIVEHYPOTHESES OF THE contrast to this, the cloacal discs of Poto- PHYLOGENY OF THE TYPHLONECTIDAE motyphlus and Typhlonectes are much A hypothesis of phylogeny for the four enlarged, the disc is recessed to varying typhlonectid genera, based on the evi- degrees, and there is variable development dence considered above, is represented by of adjacent, flap-like folds of skin. These the cladogram in Fig. 3A. Alternative phy- folds, which may be well developed in logenies to this have recently been pro- either sex, and enlarged cloacal discs gave posed. In his dendrogram, Laurent (1986) rise to the notion of "suckers" or "claspers" divided the four genera into two subgroups, in males. Additionally, there is strong sex- Chthonerpeton with Nectocaecilia and ual dimorphism in the shape of the ter- Potomotyphlus with Typhlonectes (Fig. minal shield of Potomotyphlus and Typh- 3B). Unfortunately,Laurent (1986) did not lonectes, the females having terminal provide any evidence in support of his shields that are pointed rather than bluntly phylogeny of the Gymnophiona. This lack rounded. This sexual dimorphism does not of evidence renders his hypothesis im- exist in Chthonerpeton and Nectocaecilia. mune to critical evaluation, and it remains I interpret the enlarged size of the cloacal no more than unsupported speculation. In disc, the flap-like modifications of the body terms of the evidence presented here, if terminus about the disc, and the strong Laurent's hypothesis of the phylogeny of sexual dimorphism in the shape of the ter- the Typhlonectidae were correct, it would minal shield as three shared derived fea- indicate either the independent derivation tures indicating monophyly of Potomo- of sub-triangular nares, tentacular aper- typhlus and Typhlonectes. Nussbaum tures close to the nares, relatively elongate (1979) considered "anal claspers" to be a left and right lungs, enlarged choanae, en- derived feature of Typhlonectes, and fol- larged and superficial choanal valves, in- lowing this author, Duellman and Trueb crease in the numbers of nuchal and post- (1986) erroneously considered this modi- cloacal vertebrae, and dorsally attaching fication to be typical of the Typhlonecti- fetal gills in Nectocaecilia and in the Po- dae as a whole. tomotyphlus-Typhlonectes clade, or the The morphoclines in the size of the presence of all these features in the an- choanae and the size of the choanal valves cestral typhlonectid and their subsequent provide evidence of monophyly of these multiple reversal in Chthonerpeton. two genera. Extreme enlargement of the Lescure et al. (1986) presented a new choanae and of the choanal valves are the classification of the Gymnophiona based 32 HERPETOLOGICA [Vol. 45, No. 1

Ch Ne Po Ty Ch Ne Po Ty Po Tn Tc Ch

h

- 19 f /e 9,10,11,12,13, 14,15,16,17d

- 18 A 1,2,3,4, B c ' a 5,6,7,8 FIG. 3.-Alternative hypotheses of phylogenetic relationships among typhlonectid caecilians: (A) based on evidence discussed in this paper; (B) after Laurent (1986); (C) modified from Lescure et al. (1986). Character state transformations in cladogram C (a-h) are those of Lescure et al., and dotted lines on this cladogram indicate hypothetical taxa. 1 = sub-triangular narial apertures; 2 = tentacular apertures close behind nares; 3 = left lung well developed; 4 = right lung well developed; 5 = intermediate enlarged choanal apertures; 6= intermediate enlarged choanal valves; 7 = superficial choanal valves; 8 = more than three nuchal vertebrae; 9 = several postcloacal vertebrae; 10 = extreme enlargement of choanal aperture; 11 = extreme enlargement of choanal valves; 12 = a middorsal ridge or free fold; 13 = lateral compression; 14 = fully aquatic; 15 = much enlarged cloacal discs; 16 = skin flaps bordering cloacal disc; 17 = pointed body terminus in females; 18 = dorsal attachment of fetal gills; 19 = fusion of fetal gill bases. Dashed lines indicate character states unknown in Potomotyphlus. a = II; b = III+II; c = III; d = II+III+II; e = I+Ilinc; f = III+II+I; g = II+III+II+I; h = I+II+III+II+I. Ch = Chthonerpeton; Ne = Nectocaecilia; Po = Potomotyphlus; Ty = Typhlonectes; Tn = Typhlonectes natans; Tc = Typhlonectes compressicaudus.

on the distribution of 16 character state times in the original work, it can hardly transformation series and biogeographic have been a lapsuscalami or printer'serror. data across 30 taxa representing 27 genera. Art. 32(c)(ii) of the Code clearly states that Sixty-one new suprageneric names (rather "incorrecttransliteration or latinization ... a large number considering the number of are not to be considered inadvertent taxa) were established in the new classifi- errors". Therefore Potomotyphlus is the cation. These authors established formal correct spelling. names for not only every monophyletic With regard to the 17 characters em- group, but also for every possible rank ployed by Lescure et al. (1986), the four within a clade, even when no divergence typhlonectid taxa that they consideredvary (branching) is evident. Thus, the genus Po- in only five (summarized in Table 3). Of tomotyphlus, which they hypothesized to these, one is presence of a "tail", which be the sister taxon to the rest of the typh- was scored incorrectly for Potomotyphlus lonectid groups (excluding Nectocaecilia, and probably polarized incorrectly for the which was not included in the analysis, see Typhlonectidae. A second character (fate Fig. 3C), was placed in a monogeneric epi- of the postfrontal) has the derived state family of equal rank to the group com- restricted to Potomotyphlus and is thus prising the other typhlonectids, and a fam- uninformative. These authors considered ily, subfamily, infrafamily, and tribe were < 100 vertebrae to be primitive and > 100 also named. derived. They made no attempt to justify Potomotyphlus was established by Tay- this interpretation. They scored Typhlo- lor (1968:256). The alternative spelling Po- nectes natans with the derived condition tamotyphlus appeared in the errata of and Chthonerpeton with the primitive some editions of this work. This latter spell- condition. In fact, the range of vertebrae ing was used by Lescure et al. (1986) for in Typhlonectes natans spans their arbi- this genus and for all higher taxonomic trarily defined characterstate cutoffs (Wil- rank names they proposed that utilized this kinson, unpublished), and Chthonerpeton generic name as a root. These authors pre- contains species that fall on either side of sumably considered the original spelling this character state division (Nussbaum, Potomotyphlus to be an incorrect spelling; 1986b; Nussbaum and Wilkinson, 1987; however, as this spelling appears several Taylor, 1968). Similarly problematic is March 1989] HERPETOLOGICA 33

TABLE 3.-Summary of character state distributions over the four typhlonectid taxa reported by Lescure et al. (1986). (1) Ratio of length to width: 0 = <30, 1 = between 30 and 40. (2) Number of vertebrae: 0 = <100, 1 = >100. (3) Fate of the postfrontal: 0 = fused, 1 = fused to the maxilla. (4) "Tail": 0 = present, 1 = absent. (5) Annulation. Character states represented by a 0 are primitive, those represented by a 1 are derived. Annulation character states are not polarized and are those reported by Lescure et al. (1986) in the "remarques" section of their data matrix.

1 2 3 4 5 Chthonerpeton 0 0 0 1 I+Ilinc Potomotyphlus 1 1 1 1 I+II+III+II+I Typhlonectes natans 1 1 0 0 IIinc+IIIinc+IIinc T. compressicaudus 0 0 0 0 I+Ilinc+(I) their character based on the ratio of the data. These authors preferred the clado- body length to width for which the char- gram that necessitates independent deri- acter state divisions are arbitrary, no jus- vation of two of their derived character tification of polarization is made, and the states in Potomotyphlus and in Typhlo- reported character states do not take into nectes natans (or their reversal in Chtho- account ontogenetic, seasonal, and sexual nerpeton and T. compressicaudus) over intraspecificvariation or intrageneric vari- the more parsimonious cladograms (Fig. ation, the last of which is high in Chtho- 4A,B,C) which do not require these extra nerpeton (Nussbaum, 1986b; Taylor, steps. In doing this, Lescure et al. seem to 1968). have ignored some of their own "evi- The final differences involve annula- dence" and relied solely on their interpre- tion. Whereas some of the other characters tation of annular evolution to determine employed by Lescure et al. (1986) were the genealogy of the typhlonectid taxa. In- adequately defined, in the case of trunk deed, the only instances of character evo- annulation, both the characterstates them- lution mapped onto the typhlonectid por- selves and their polarities were not made tion of their cladogramconcern annulation. clear. In fact, the reported character states Lescure et al. preferred to rely on annu- are completely at odds with presently ac- lation rather than use any of their other cepted concepts and observations.Lescure characters despite their own caveats that et al. (1986) referred to another of their the annulation states of typhlonectids are papers, Renous et al. (1986), for a discus- difficult to determine and that those they sion and justification of the morphoclines reported are not certain. Over-reliance on that they employed. However, Renous et the annulation character is also apparent al. (1986) did not, as they claimed, provide in their treatment of the majority of non- these justificationsfor the majorityof their typhlonectid taxa. morphoclines. In their discussion of the Based on the paraphyly of Typhlonectes evolution of annulation, they asserted that indicated in their hypothesis, Lescure et secondary and tertiary annuli form ante- al. (1986) established a new genus, Pseu- riorly first,respecting a cephalo-caudal on- dotyphlonectes, to receive T. natans. The togenetic gradient. Data from ontogenetic supposed paraphyletic status of Typhlo- series of a number of caecilian species in- nectes restssolely on the position of Chtho- dicate that this assertion is erroneous and nerpeton relative to the two species of that Lescure et al. almost certainly have Typhlonectes that were considered. That an erroneous notion of the evolution of this position is erroneous is indicated by caecilian annulation (Nussbaum and Wil- all the evidence that I have discussed. kinson, in press). In terms of the evidence that I have I have reanalyzed the data from Lescure presented, the hypothesis of Lescure et al. et al. (1986) for the Typhlonectidae. The (1986), if correct, would necessitatethe loss results of this reanalysis (summarized in in Chthonerpeton of sub-triangularnares, Fig. 4) show that their published clado- tentacular aperturesclose to the nares, rel- gram is not the one that best fits their own atively elongate lungs, enlarged choanae, 34 HERPETOLOGICA [Vol. 45, No. 1

Ch Tc Tn Po Ch Tc Tn Po

4,5 X/ \ 4 ,4,5b 4,5a 5d 3,4,5b

5d \ f 1,2 \ \ f 1,2

A ~~~~~~~~~B tC5c t5c

Tc Ch Tn Po Po Tn Tc Ch

5 5a 4r 3,5b 3,4,5 5a

\ X f 1~~~~,2 r\ \ r 2 r,5d

C 7Zc D H125c

FIG. 4.-Reanalysis of the data of Lescure et al. (1986), showing the distribution of the character states summarized in Table 3. Cladograms A, B, and C are three equally most parsimonious interpretations of the data (all with a consistency index of 0.875). Cladogram D is the topology published by Lescure et al. (1986) with character state changes plotted so as to minimize the number of character state changes (consistency index of 0.7). Annulation states: a = I+Ilinc; b =I+II+III+Il+l; c = IIinc+IIIinc+Ilinc; d = I+IIinc+(I). Ch = Chthonerpeton; Po = Potomotyphlus; Tn = Typhlonectes natans; Tc = Typhlonectes compressicaudus. For each cladogram, where homoplasy is indicated, alternative equally parsimonious interpretations of character evolution (convergence or reversal), which do not affect tree topology, are possible. Only one interpretation is shown. Non-annulation character state changes are from primitive to derived except where indicated. r = reversal. enlarged and superficial choanal valves, hypothesis that it was used to support is lateral compression, middorsal ridges or contradicted by all the evidence discussed free folds, more than three nuchal and sev- here. eral postcloacal vertebrae, modified ter- If my interpretation of typhlonectid minal shields, enlarged cloacal discs, dor- phylogeny is correct, then not only is the sally attaching gills with fused bases, and genus Pseudotyphlonectes invalid, but so a fully aquatic lifestyle. are all the new higher taxonomic cate- The numerous taxa erected by Lescure gories that Lescure et al. (1986) applied to et al. (1986) within their "Potamotyphloi- the Typhlonectidae (sensu Taylor, 1968), dae" were each provided with a diagnosis. including the new family Potamotyphli- Which characters were employed for dae. I consider that Pseudotyphlonectes which diagnosis might at first seem arbi- Lescure, Renous, and Gasc is a junior syn- trary as the whole epifamily is monoge- onym of Typhlonectes Peters. Further- neric. In fact, successively lower taxonom- more, because the character of the annu- ic ranks were diagnosed through the lation of the trunk as used by Lescure et exclusion of taxa not known to exist, but al. has resulted in an unsatisfactory hy- which are hypothesized as intermediate pothesis of relationships within the Typh- stages in the evolution of the trunk an- lonectidae, I have no confidence in its sim- nulation. Considerabledoubt must now be ilar application elsewhere and consider the cast on the validity of Lescure et al.'s in- entire classification of the Gymnophiona terpretation of the trunk annulation char- proposed by Lescure et al. to be unsoundly acter, because in addition to the anomalous based. characterstates reported, the phylogenetic Taylor (1968:40) commented that "Un- March 1989] HERPETOLOGICA 35

til greater study is focussed on the cranium drew Stimson provided advice on taxonomic and no- and other parts of the skeleton and also on menclatural issues. Ron Nussbaum and David Bay helped prepare the figures and Susan Morseau helped the viscera, it would seem rather futile to prepare the manuscript. I thank the editors of Her- try and trace evolutionary relationshipsin petologica for their comments and advice. This work detail". Though much ground has been would not have been possible without the financial gained in our knowledge of caecilian phy- assistance of Joanne Skerratt to whom I am especially logeny, we have not come so far down the grateful. road from 1968 that Taylor's cautionary words are without significance. While I do LITERATURE CITED not want to discourage attempts to recon- BARRIO, A. 1969. Observaciones sobre Chthoner- struct the phylogeny of the Gymnophiona, peton indistinctum (Gymnophiona, Caeciliidae) y it is recommended that such attempts be su reproduccion. Physis 28(77):499-503. decoupled from taxonomic treatment until DUELLMAN, W. E., AND L. TRUEB. 1986. Biology a greater understanding is achieved. The of . McGraw-Hill, New York. ELTER, 0. 1981. Cataloghi V-La collezione er- alternative to this is a period of great taxo- petologica del Museo di Zoologia dell'Universita di nomic instability and a plethora of names Torino. University of Torino. and ranks many of which will prove to be GAYMER, R. 1971. New method of locomotion in of no utility whatsoever. limbless terrestrial vertebrates. Nature 234:150-151. LAURENT, R. F. 1986. Evolution des gymnophiones. Acknowledgments.-I thank the following indi- Pp. 606-608. In P. P. Grasse (Ed.), Traite de Zoo- viduals and institutions for the loan of or access to logie 14. specimens used in this study: C. W. Myers,American LESCURE, J., S. RENOUS, AND J. P. GASC. 1986. Pro- Museumof Natural History (AMNH);E. V. Malnate, position d'une nouvelle classification des amphi- Academy of Natural Sciences, Philadelphia (ANSP); biens gymnophiones. Mem. Soc. Zool. France 43: E. N. Arnoldand B. T. Clarke,British Museum (Nat- 145-177. ural History) (BMNH); R. C. Drewes, California MURPHY, J. B., H. QUINN, AND J. A. CAMPBELL. 1977. Academy of Sciences (CAS);C. J. McCoy, Carnegie Observations on the breeding habits of the aquatic Museum (CM); the curators, Estacion Biologia de caecilian Typhlonectes compressicaudus. Copeia Rancho Grande (EBRG);R. F. Laurent, Fundacion 1977:66-69. Miguel Lillo, Tucumen (FML); R. F. Inger, Field NAYLOR, B. G., AND R. A. NUSSBAUM. 1980. The Museum of Natural History (FMNH); W. E. Duell- trunk musculature of caecilians (Amphibia: Gym- man, University of Kansas,Museum of Natural His- nophiona). J. Morphol. 166:259-273. tory (KU); J. Wright, Los Angeles County Museum NUSSBAUM, R. A. 1977. : A new of Natural History (LACM);J.-M. Exbrayat and M. family of caecilians (Amphibia: Gymnophiona). Delsol, Faculte Catholique des Sciences de Lyon Occ. Pap. Mus. Zool. Univ. Michigan 682:1-30. (LYON); the curators, Universidad Central, Museo 1979. The taxonomic status of the caecilian de Biologia, Caracas (MBUCV); P. Alberch and J. genus Uraeotyphlus Peters. Occ. Pap. Mus. Zool. Rosado, Museum of Comparative Zoology, Harvard Univ. Michigan 687:1-20. (MCZ);V. Mahnertand J.-L. Perret, Museum d'His- . 1985. Systematics of the caecilians (Am- toire Naturelle de Geneve (MHNG);J. Lescure, Mu- phibia: Gymnophiona) of the family Scolecomor- seum National d'Histoire Naturelle, Paris (MNHN); phidae. Occ. Pap. Mus. Zool. Univ. Michigan 713: G. Lenglet, Institut Royal des Sciences Naturelles de 1-49. Belgique (MRHN); F. Tiedemann and A. Cabella, 1986a. The taxonomic status of Lutkeno- Naturhistorisches Museum, Wien (NMW); R. A. typhlus brasiliensis (Lutken) and Siphonops con- Nussbaum, University of Michigan, Museum of Zo- fusionus Taylor (Gymnophiona: Caeciliidae). J. ology (UMMZ);W. Bohme, ZoologischesForschung- Herpetol. 20:441-444. sinstitut und Museum AlexanderKoenig (ZFMK);L. 1986b. Chthonerpeton onorei, a new cae- J. Borkin, ZoologicalInstitute, Academy of Sciences, cilian (Amphibia: Gymnophiona: Typhlonectidae) Leningrad(ZIN); R. Gunther,Universitat Humboldt, from Equador. Revue Suisse Zool. 93:911-918. Zoologisches Museum, Berlin (ZMB); H. W. In press. On the status of Copeotyphlinus Koepke,H. Wilkens,and R. Dohse, UniversitatHam- syntremus, Gymnopis oligozona, and Minascae- burg, Zoologisches Museum (ZMH); J. Rasmussen, cilia sartoria (Gymnophiona, Caeciliidae): A com- ZoologiskMuseum, Copenhagen (ZMUC); U. Gruber, edy of errors. Copeia. ZoologischesStaatssammlung, Munchen (ZSM). NUSSBAUM, R. A., AND C. GANS. 1981. On the I am indebted to Orsetta Elter and her staff at the Ichthyophis (Amphibia: Gymnophiona) of Sri Lan- Museo di Zoologia,Torino for facilitating my visit to ka. Spolia Zeylanica 35:137-154. that Institution and for the numerous courtesies ex- NUSSBAUM, R. A., AND M. WILKINSON. 1987. Two tended to me there. Lilia Capocaccia and Giuliano new Chthonerpeton (Amphibia: Gymnophiona: Doria made me welcome at the Museo Civico di Typhlonectidae) from Brazil. Occ. Pap. Mus. Zool. Storia Naturale, Genova. Ron Nussbaum provided Univ. Michigan 716:1-15. indispensible encouragement and discussions. An- In press. On the classification and phylog- 36 HERPETOLOGICA [Vol. 45, No. 1

eny of caecilians (Amphibia: Gymnophiona), a crit- (lectotype); AMNH A25506, A23508; CM 65022-25, ical review. Herpetol. Monogr. 65037; CAS 85521; FML 032284; KU 197262-64; RENOUS,S., J. LESCURE, AND J.-P. GASC. 1986. Der- BMNH 83.1.19.2, 89.8.24.4, 1926.5.29.7; IMZT An mal rings and vertebral segments in the Gymno- 786; MNHN 4299; NMW 9148; USNM 65538, 95863, phiona. Pp. 297-302. In Z. Rocek (Ed.), Studies in 257179; ZIN 1795.1, 1793, 1382, 1759; ZMB 6807, Herpetology. Charles University, Prague. 34527-28; ZMH A00256. C. onorei: MNHG 2251.06 SAVAGE, J. M., AND M. H. WAKE. 1972. Geographic (holotype), 2251.07 (paratype). C. viviparum:BMNH variation and systematics of the middle American 1947.2.13.84 (holotype); MCZ 24593; ZIN 1762.1; caecilians, genera Dermophis and Gymnopis. Co- ZFMK 27687. Nectocaecilia petersii: BMNH 61.9.2.6 peia 1972:680-695. (holotype); AMNH 100590-91, 100890; MBUCV 960- SWOFFORD, D. L. 1985. PAUP: Phylogenetic Anal- 61. N. ladigesi: ZSM 245 (holotype). Potomotyphlus ysis Using Parsimony. Version 2.4. Illinois Natural kaupii: ZFMK 27684 (holotype); ZMH A00259-63, History Survey, Champaign, Illinois. A01017-18; CAS 10848; CM 2906,2908; ANSP 4926- TANNER, K. 1971. Notizen zur pflege und zum ver- 27; MCZ A96785; BMNH 98.10.17.7; FMNH 206189; halten einiger blindwiihlen (Amphibia: Gymno- KU 128088; USNM 101105, 166415,305344. P. mela- phiona). Salamandra 7:91-100. nochrous: NMW 9147 (holotype). Typhlonectes an- TAYLOR, E. H. 1968. The Caecilians of the World: guillaformis: AMNH 56252 (holotype). T. compres- A Taxonomic Review. University of Kansas Press, sicaudus: MNHN 4269 (lectotype); CM 90090-91; Lawrence. CAS 125421-23; UMMZ 82854; LYON 81.1.1-4, 1970. An aquatic caecilian from the Mag- 81.1.7-15, 81.1.17-32; MRHN 8681-83, 8743; BMNH dalena River, Colombia, South America. Univ. 1982.103-104, 1916.4.12.1-2; ZFMK 42781; USNM Kansas Sci. Bull. 48:845-848. 201693, 201695; MCZ A85381. T. eiselti: NMW 9146 WILKINSON, M. 1988. On the statusof Nectocaecilia (holotype). T. natans: ZMB 9522 (cotype); LACM cooperi Taylor, with comments on the genus Nec- 67414-517; UMMZ 172649-50; CAS 153871; IMZT tocaecilia Taylor (Amphibia: Gymnophiona: Ty- An-781; MSNG 379/A. phlonectidae). J. Herpetol. 22:119-121. Outgroups.-Caecilia gracilis:UMMZ 4710, 4711, 52507, 76676. C. nigricans: UMMZ 121035, 124109. Accepted: 22 February 1988 C. pachynema: UMMZ 82902-03. C. tentaculata: Associate Editor: John Iverson UMMZ 89459, 177895. Dermophis mexicanus: UMMZ 152768-75. Epicrionops marmoratus: BMNH APPENDIX I 1956.1.15.87 (holotype). E. petersi petersi: UMMZ 163243 (paratopotype). Gymnopis multiplicata: Material Examined UMMZ 142682, 13170-72. Lutkenotyphlus brazil- Typhlonectidae. -Chthonerpeton braestrupi: iensis: UMMZ182015. Microcaecilia albiceps: UMMZ ZMUC R0234 (holotype). C. corrugatum: ZMH 83051. M. unicolor: UMMZ 173394. Oscaecilia och- A00235 (holotype); ANSP 13948 (paratype). C. eru- rocephala:UMMZ 167596-99. Siphonopsannulatus: gatum: ZMUC R0238 (holotype). C. helmichi: ZSZM IMZT An-782-784; UMMZ 89460, 177895. S. pau- 1/1964 (holotype). C. indistinctum: ZMUC R0235 lensis: UMMZ 109665, 137582-83, 187585.