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1 AMPHIBIA: : Potomotyphlus, Potomotyphlus kaupii Catalogue of American and DESCRIPTION. Potomotyphlus kaupii is Reptiles 924 an aquatic endemic to much of northern South America. The longest known Ledesma, D. T. 2019. Potomotyphlus, specimen of Potomotyphlus kaupii has a to- Potomotyphlus kaupii. tal length of 695 mm (Dunn 1942; Taylor 1968); total lengths for additional specimens Potomotyphlus Taylor range from 33–555 mm (Maciel and Hoog- Cecilia de Río (River Caecilian) moed 2011). Along the body, the number of primary annuli ranges 83–102. In the orig- Potomotyphlus Taylor 1968:11, 256. Type spe- inal description of the , 104 primary cies: “ kaupii (Berthold),” by des- annuli were reported (Berthold 1859), how- ignation. ever, that count also may have included the Potomotyphlops: Taylor 1968:38. Lapsus (see collars (Taylor 1968). Nuchal grooves are po- Nomenclatural History). sitioned laterally and ventrally on the body Potamotyphlus: Taylor 1969a:600. Lapsus (see (Maciel and Hoogmoed 2011). The primary Nomenclatural History). annuli are interrupted dorsally and may not Potomotuphus: Ledesma 2019. Lapsus. be well defined by distinct grooves (seeRe - Potomotyplus: Ledesma 2019. Lapsus. marks). There are no secondary annuli pres- ent; however, pseudosecondaries may be vis- CONTENT. One species, Potomotyphlus ible (Nussbaum and Wilkinson 1989). The kaupii, is recognized (see Nomenclatural last three to six annuli are laterally distinct history). and no annuli are present from the approxi-

Figure 1. Body of Potomotyphlus kaupii (Museum of Vertebrate Zoology [MVZ] 173770) showing rela- tive size of head and collar compared to body. Scale bar is 1 cm. Photographed by Simon Scarpetta. 2

10°N 70°W 60°W 50°W

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Map. Geographic distribution of Potomotyphlus kaupii. The open circle represents the type locality, the question marks indicate uncertainty in the reported locality, and the black dots represent localities from the literature corresponding to the following: 1. Rio Santiago, La Poza, Peru (Catenazzi and Venegas 2012); 2. Pucayacú, Pastaza Province, Ecuador (Taylor and Peters 1974); 3?. Atrato River System, Colombia (Taylor 1968) (see Geographic Distribution); 4. Rio Guayabero, La Macarena, Meta, Colombia (Lynch 2000); 5. Rio Meta, Puerto Lòpez, Colombia (Lynch 2000); 6. Rio Itaya, San Antonio, Iquitos, Peru (Taylor 1968); 7. Rio Ampiyac, Iquitos, Peru (Taylor 1968); 8. Rio Caquetà, La Pedera, Amazonas, Colombia (Lynch 2000); 9. Rio Branco, Acre, Brazil (Maciel and Hoogmoed 2011); 10. Madeira River, municipality of Porto Velho, Rondônia, Brazil (Oliveira et al. 2012); 11. Cuidad Bolivar, Venezuela (Berthold 1859); 12. Cachoeira do Bem Querer, Roraima, Brazil (Maciel and Hoogmoed 2011); 13. Ilha Marchantaria, Amazonas, Brazil (Maciel and Hoogmoed 2011); 14. UHE Balbina, Presidente Figueiredo, Amazonas, Brazil (Maciel and Hoogmoed 2011); 15?. Mato Grosso, Brazil ( microcephalum) (A. Miranda-Ribeiro 1937) (see Geographic Distribution); 16. Apiacás River, Mato Groso, Brazil (Alves-Silva et al. 2017); 17. Cachoeira Porteira, Oriximiná, Pará, Brazil (Maciel and Hoogmoed 2011); 18. Sanarém, Pará, Brazil (Maciel and Hoogmoed 2011); 19. “Cayenne”, (Taylor 1968); 20. Oyapock River, Saint-Georges-de-l’Oyapock, French Guiana (Marty et al. 2007a); 21. Rio Amazonas, municipality of Macapá, Amapá, Brazil (Caramaschi and Pombal 2000); 22. Rio Xingu, Melgaço, Pará, Brazil (Maciel and Hoogmoed 2011); 23. Abaetetuba, Pará, Brazil (Maciel and Hoogmoed 2011); 24. Belém, Pará, Brazil (Maciel and Hoogmoed 2011); 25. Acará, Pará, Brazil (Maciel and Hoogmoed 2011); 26. Tucuruí, Pará, Brazil (Maciel and Hoogmoed 2011); 27. Jacundá, Pará, Brazil (Maciel and Hoogmoed 2011); 28. Serra dos Carajás, Pará, Brazil (Maciel and Hoogmoed 2011); 29. Tocantins River, Estreito hydroelectric powerplant, Maranhão, Brazil (Cintra et al. 2010); 30. Usina Hidroeletrica Serra da Mesa, Goiás, Brazil (Maciel and Hoogmoed 2011). Map was created in part using ArcMap™ © Esri. Layers used include “World Countries (Generalized)” source Esri and “South American Rivers” source user madalynbotkin. Account 924 3

Figure 2. Head of Potomotyphlus kaupii (MVZ 173770) in lateral view. Scale bar is 5 mm. Photo by Simon Scarpetta. mate level of the vent to the terminus (Maciel posterior portion of the body (see Remarks). and Hoogmoed 2011). The head and collars Additionally, some specimens have a ventral of the are noticeably small, making ridge that extends approximately 30 mm in them narrower in comparison with the por- front of the cloacal disk (Wilkinson and Nuss- tion of the body immediately posterior to baum 1997). Dermal and subdermal scales the collars (Taylor 1968). The skull is slightly are absent and paired anal papillae are vari- compressed dorsoventrally. A small specimen ably present. In one study, paired anal papillae of Potomotyphlus kaupii (Carnegie Museum were present on two of the 57 male specimens [CM] 2906) has head and body proportions examined (Maciel and Hoogmoed 2011). The that are the same as other typhlonectids of vent disk is extended anteriorly and rounded similar size, demonstrating ontogenetic vari- posteriorly with a few anteromedial dentic- ation in head size (Wilkinson and Nussbaum ulations that may be elongated (Maciel and 1999). The tongue is completely attached to Hoogmoed 2011). The vent of males has been the mandibular mucosa anteriorly. There are hypothesized to serve as a clasper used during two large narial plugs. The snout projects copulation (Duellman and Trueb 1986, 1994; anteriorly beyond the mouth. The eye is not Taylor 1968; Taylor and Peters 1974; but see covered by bone and is usually visible (see Wilkinson 1989). The vent disk is surrounded Remarks) (Maciel and Hoogmoed 2011). by fleshy folds of skin in both sexes and there There is intraspecific variation in whether the is variation in whether the disk is depressed large, sub-triangular nares are visible when (Maciel and Hoogmoed 2011). Sexual dimor- viewed dorsally. A small anterior tentacle is phism and ontogenetic variation are present located below the nostrils, near the margin of in the vent region with males having a more the mouth, and cannot be seen dorsally (Ma- sunken disk than females; smaller individu- ciel and Hoogmoed 2011). als lack a depression. The fleshy skin flap -sur The body is laterally compressed and rounding the cloacal disk may obscure the possesses a dorsal free fold of skin (Wilkin- disk from view in larger female specimens. son 1988) that is especially prominent on the The shape of the terminus is dimorphic with 4 females having a narrower and more pointed In life, specimens of Potomotyphlus terminus than males (Maciel and Hoogmoed kaupii are described as grayish or brownish 2011). The number of anal denticulations in color with darker annular grooves (Ma- range from 7–11 with a similar number at the ciel and Hoodmoed 2011; Tapley et al. 2019). posterior and anterior edge of the vent (Ma- A recently deceased female individual was ciel and Hoogmoed 2011). Potomotyphlus described as being blue-grey dorsally and kaupii has a relatively large number of teeth white-blue ventrally with a pale head, though that are tightly packed on relatively large once placed in alcohol the blue dorsal surface dentigerous areas of tooth bearing bones turned dark brown and the white-blue ven- (Wilkinson and Nussbaum 1997). There are tral surface turned grey (Marty et al. 2007a). four series of monocuspid teeth. In total, A photograph of an uncollected specimen the maximum premaxillary-maxillary tooth shows the color of a live individual as being count is 60 with no distinct variation in size. white ventrally with a gray dorsal coloring Prevomerine-palatine teeth have a maximum that transitions into a blue color on the head count of 53 and are smaller than the pre- (Marty et al. 2007b). maxillary-maxillary teeth. The dentary teeth Preserved specimens of Potomotyphlus number up to 64 and are approximately the kaupii were described as being similar to same size as the premaxillary-maxillary teeth; the blue-gray of natans and however, the posterior dentary teeth are rela- darker color of Typhlonectes compressicauda tively smaller in size than the anterior teeth. (Wilkinson and Nussbaum 1997). The color Splenial teeth have a maximum number of of the annular grooves of some individuals 12 and are approximately the same size as of Potomotyphlus kaupii was described as be- the prevomerine-palatine teeth (Maciel and ing similar to the purple color of preserved Hoogmoed 2011). Variation in the number specimens of (Wilkinson and of teeth was observed to be a function of size Nussbaum 1997). The color of preserved Po- with larger specimens having more teeth than tomotyphlus kaupii has also been described as smaller ones (Taylor 1968). being dorsally gray-olive with the sides and

Figure 3. Head of Potomotyphlus kaupii (MVZ 173770) in ventral view. Scale bar is 5 mm. Photo by Simon Scarpetta. Account 924 5

Figure 4. Terminus of Potomotyphlus kaupii (MVZ 173770) from ventral view showing cloacal disk. Scale bar is 5 mm. Photo by Simon Scarpetta. venter having a yellowish-olive color with a and Typhlonectes and more anterior to the eye cream-colored vent (Taylor 1968). The ho- compared to Chthonerpeton (Wilkinson and lotype for the now-synonymized species Po- Nussbaum 1997). Potomotyphlus is unique tomotyphlus melanochrus, Naturhistorisches among typhlonectids in having partially Museum, Wien [NMW] 9147, was described fused choanal valves with a concealed aper- as being black throughout except for a lighter ture and differs fromAtretochoana in having area around the eyes (Taylor 1968). a line of fusion of the valves marked with a thickened ridge (Wilkinson and Nussbaum DIAGNOSIS. Potomotyphlus kaupii occurs 1999). Within Typhlonectidae, the monotyp- sympatrically with other aquatic ic genera Potomotyphlus and including Typhlonectes compressicauda in have the greatest number of tooth positions Colombia, French Guiana, and Brazil (Maciel on the dentary and the teeth of Potomotyphlus and Hoogmoed 2011) and with Atretochoana have a more conical shaft, narrow base, and a eiselti in the Mosqueiro region and possibly well-developed lateral flange (Wilkinson and the Cachoeira Santo Antônio area of Brazil Nussbaum 1999). The orientation and relative (Hoogmoed et al. 2011). The shape of the size of the nostrils, anteriorly directed and vent (cloacal) disk in Potomotyphlus kaupii, large in Atretochoana eiselti but not in Poto- elongate and narrow anteriorly while round- motyphlus kaupii, and the relatively flatter ed posteriorly, is unique among typhlonectids and wider head of Atretochoana eiselti com- (Wilkinson et al. 2011). In adult specimens pared to Potomotyphlus kaupii can be used to of Potomotyphlus kaupii, the anterior region distinguish the two species (Hoogmoed et al. of the body, including the head and collars, 2011). is disproportionately small compared to the region just posterior, a feature that distin- PHYLOGENETIC RELATIONSHIPS, guishes the species from other caecilians PUBLISHED DESCRIPTIONS, ILLUS- (however see Remarks). The tentacular ap- TRATIONS, DISTRIBUTION, FOSSIL RE- erture of Potomotyphlus is located more pos- CORD, and PERTINENT LITERATURE. terior to the nares compared to Nectocaecilia See species account. 6

Figure 5. Body of Potomotyphlus kaupii showing a dorsal free fold. Live individual from the Sedgwick County Zoo, Kansas. Photo by Charles Withnell.

NOMENCLATURAL HISTORY. Both the and the classification was not adopted by generic name and the species were described subsequent authors, with the exception of the by Taylor (1968:256 and 1968:257, respective- use of Potomotyphlidae (Exbrayat 2000, Les- ly). The species Potomotyphlus melanochrus cure and Renous 1988, Pefaur 1992, Renous was also described in 1968 (Taylor 1968:263) and Gasc 1989) and the mention of Potamo- but was subsequently synonymized with Po- typhloidea (Frost et al. 2006). An argument tomotyphlus kaupii by Nussbaum and Wilkin- was made to regard the original spelling “Po- son (1989). tomotyphlus” (Taylor) as the correct spelling The spelling Potamotyphlus“ ” was used by (Wilkinson 1989) because Article 32.5.1. of Taylor (1969a, 1969b) in the place of the orig- the International Code of Zoological No- inally assigned name by Taylor (1968), Poto- menclature states that, “Incorrect translitera- motyphlus. The spellings used by Lescure et tion or latinization, or use of an inappropriate al. (1986) followed the spelling used by Tay- connecting vowel, are not to be considered lor (1969a, 1969b) in that the root, “Potamo,” inadvertent errors” (International Commis- appeared in the proposed epifamily “Potamo- sion on Zoological Nomenclature 1999:39). typhloidea,” family “Potamotyphlidae,” sub- In other publications, however, the generic family “Potamotyphlinae,” and infrafamily name Potamotyphlus was regarded as the cor- “Potamotyphlilae.” However, the tribe pro- rect spelling because that spelling appeared posed by Lescure et al. (1986), “Potomotyph- in the errata included in some editions of lini,” follows the root of the original spelling Taylor (1968) (e.g., Frost 2018). Additionally, of Potomotyphlus (Taylor). The the name “Typhlonectus kaupii” was used by proposed by Lescure et al. (1986) was critical- Frost (2018) to reference a spelling purported ly reviewed (Nussbaum and Wilkinson 1989) to appear in Gines (1959), however, I can only Account 924 7 find Typhlonectes“ kaupii” in Gines (1959:98) Ciudad Bolivar, Venezuela]. Holotype [by rather than the name used by Frost (2018). implication, Böhme and Bischoff 1984], REMARKS. The annuli ofPotomotyphlus Zoological Research Institute and Mu- kaupii are often not defined with distinct seum Alexander Koenig [ZFMK] 27684, grooves but may instead be marked by line collector and date of collection unknown. glands and pigmentation that may be ho- Not examined by author. mologous to the grooves in other caecilians kaupii: Keferstein 1867:361. (Wilkinson and Nussbaum 1999; but see Caecilia dorsalis Peters 1877:459. Type local- Nussbaum and Wilkinson 1989). ity “Angostura (Ciudad Dolivar {sic}) am The visibility of the eye was reported to Orinoco” [=Ciudad Bolivar, Venezuela in vary ontogenetically so that small specimens the Orinoco]. Holotype, Zoological Mu- (100–200 mm in length) have a more visible seum Berlin [ZMB] 9092 (Bauer et al. eye compared to the eyes of larger specimens, 1993; see Remarks), juvenile, collected which may be obscured or covered by skin by Dr. Sachs, date of collection unknown. (Taylor 1968). However, the visibility of the Not examined by author. eye did not correlate with body size and in Typhlonectes dorsalis: Peters 1880:941. fact the eye was visible in a majority of spec- Typhlonectes kaupii: Boulenger 1891:457. imens examined by Maciel and Hoogmoed Recognized synonymy of Typhlonectes (2011). dorsalis and Siphonops kaupii. The use of the terms 'free fold' and 'ridge' Thyphlonectes dorsalis: Fuhrmann 1914:124. follows definitions provided by Wilkinson Lapsus. (1988). Additionally, the distinctiveness of a Chthonerpeton microcephalum A. de Miran- free dorsal fold of skin may vary between live da-Ribeiro 1937:64. No type given, only and preserved specimens (Wilkinson 1988). type locality “Matto-Grosso” (see Geo- The apomorphic condition found inPoto - graphic Distribution). Holotype lat- motyphlus kaupii of the head and collars be- er designated by P. de Miranda-Ribeiro ing disproportionately small compared to the (1955): Museu Nacional, Rio de Janeiro, body was reported in the holotype of Dermo- “M. N. n.° 539,” no date of collection giv- phis sertentrionalis (Taylor 1968) (synonym en. Collected by Rondon (Dunn 1942 ) or of mexicanus) and may represent “Comissão Rondon” (Bokermann 1966). a teratology in that holotype (Wilkinson and Not examined by author. Nussbaum 1999). Chthonefpton microcephalum: Travassos 1955:XXXV. Lapsus. ETYMOLOGY. The namePotomotyphlus Chthonespton microcephalum: P. de Miran- appears to be derived from the Greek word da-Ribeiro 1955:392. Lapsus. (see Re- potamos meaning river and the Greek word marks). tuphlos meaning blind. The reference to a riv- Potomotyphlus kaupii Taylor 1968:11, 257. er seems to be connected to the aquatic life- Type not specified (but seeRemarks). style of the species and the reference to the Potomotyphlus melanochrus Taylor 1968:263. word blind may be related to the origin of the Type locality “Brazil.” Holotype, Naturhis- name “caecilian” from the Latin word caecus torisches Museum, Wien, Austria [NMW] meaning blind. 9147. Collected in 1875 by unknown col- lector, additional tag number “II. 120” Potomotyphlus kaupii Berthold (Gemel et al. 2019). Not examined by au- Kaup's Caecilian thor. Potomotyphlus kaupi: Frank and Ramus Caecilia kaupii Berthold 1859:181. Type lo- 1995:26. Lapsus. (see Nomenclatural cality “ex Angostura” [from Angostura] [= History). 8

CONTENT. There are no recognized subspe- by Kerferstein (1867). Descriptions of the ex- cies. ternal morphology were provided by Peters (1877, 1880). A description of Potomotyphlus DESCRIPTION and DIAGNOSIS. See ge- kaupii, under the generic name Typhlonectes, neric account. was also published by Nieden (1913, 1965), and a description of the morphology of the PHYLOGENETIC RELATIONSHIPS. internal organs, skin, and sagittal section of Potomotyphlus kaupii is placed within the the head was provided by Fuhrmann (1914). family Typhlonectidae (Taylor 1968) or, de- Descriptions were also published by A. de pending on the classification, within the sub- Miranda-Ribeiro (1937, 1955), and Dunn family Typhlonectinae (Frost et al. 2006). The (1942). A diagnosis and description of Po- species is monotypic within the epifamily tomotyphlus kaupii was provided by Taylor Potamotyphloidea and numerous lower clas- (1968). A description of the cranial osteology sifications therein proposed by Lescure et of the species was provided by Taylor (1969a) al. (1986). That classification was not widely and a brief overview of the osteology of Poto- adopted by other authors and the methods motyphlus was given by Wake (2003). A de- used in the study were rejected by Nuss- scription of the single specimen known from baum and Wilkinson (1989) and Wilkinson Ecuador was provided by Taylor and Peters (1989). Phylogenetic analyses including Po- (1974). Descriptions of trunk musculature of tomotyphlus kaupii, based on morphological caecilians, including Potomotyphlus kaupii, (Wilkinson and Nussbaum 1999) and molec- were published by Nussbaum and Naylor ular (Maciel et al. 2016) data, recovered the (1982). Comprehensive descriptions of the species as part of a clade consisting of aquatic osteology, myology, circulatory and respira- and semi-aquatic typhlonectids including the tory morphology with comparisons to other genera Atretochoana, Chthonerpeton, Necto- typhlonectids were provided by Wilkinson caecilia, Potomotyphlus, and Typhlonectes. Po- and Nussbaum (1997, 1999) and an abstract tomotyphlus kaupii is the monotypic species on histological examination of the skin was for the genus and was hypothesized, based on published by Canepa et al. (1999). A descrip- shared morphology, to be the sister taxon of tion with comments on the natural histo- Typhlonectes (Wilkinson 1989). After the de- ry was provided by Marty et al. (2007a) and scription of the monotypic genus Atretochoa- comments on the coloring of a live individual na (containing the lungless species Atretocho- were given by Marty et al. (2007b). A mor- ana eiselti), Potomotyphlus and Atretochana phological diagnosis of Potomotyphlus was were recovered as sister taxa following an ex- given by Wilkinson and Nussbaum (2006) tensive morphological phylogenetic analysis and Wilkinson et al. (2011). A description (Wilkinson and Nussbaum 1999). However, and diagnosis were given by Maciel (2009) a subsequent molecular phylogenetic analy- and Maciel and Hoogmoed (2011). sis recovered Potomotyphlus kaupii as more closely related to species within the genus ILLUSTRATIONS. Color photographs Typhlonectes, suggesting that the shared mor- were published by Alves-Silva et al. (2017), phological characters of Potomotyphlus and Marty et al. (2007b), Oliveira et al. (2012), Atretochoana may have a more complex his- Pough et al. (2004), and Ruiz-Carranza et al. tory than previously hypothesized (Maciel et (1996). Black-and-white photographs of the al. 2016) (see Remarks). skull were published by Sherratt et al. (2014) and Taylor (1969a, 1969b) (see Remarks). PUBLISHED DESCRIPTIONS. The species Black-and-white photographs of head and was originally described by Berthold (1859) body were provided by Hillman et al. (2009), and an additional description was provided Marty et al. (2007a), Pough et al. (1998, Account 924 9

2001), Taylor (1968), and Wilkinson (1989). the state of Rondônia) (Bokermann 1966). Black-and-white illustrations of the head, Furthermore, in a correspondence reported terminus, and body were published by Peters by Dunn (1942:539), Dr. Joseph Bailey stat- (1880). Illustrations of the head, body, and ed that the Mato Grosso collections “all came sagittal cross section of the head were pub- from the northern and western sections of lished by Fuhrmann (1914) and illustrations the state” and it is likely that “most of it came of the head and terminal regions were pub- from the Serra de Parecis or along what is lished by Taylor (1968). Black-and-white di- now Rio Roosevelt.” Specimens have since agrammatic illustrations of the cloacal disk, been reported from Mato Grosso (Alves-Sil- choanal valves, cranial bones, and lower jaws va et al. 2017) and from Rondônia (Oliveira et were presented by Wilkinson and Nussbaum al. 2012) confirming the presence of the spe- (1997), illustrations of the nasopremaxilla, cies in those areas. A specimen, Institut Royal os basale, choanae, basibranchials, cerato- des Sciences Naturelles de Belgique (IRSNB) branchials, arytenoids, and cranial muscles 4544, was reported from “Cayenne”, French were published by Wilkinson and Nussbaum Guiana, but the accuracy of that locality was (1999), and an illustration of a right anteri- questioned, and the species was not includ- or dentary tooth was presented by Wilkinson ed in checklists for French Guiana (Lescure (1991). A black-and-white scanning elec- and Marty 2000). The subsequent discovery tron micrograph of anterior dentary teeth of a specimen in French Guiana (Marty et al. was published by Wilkinson and Naussbaum 2007a) confirmed the presence of the species (1997). An x-ray photograph was published in that area. A specimen, Academy of Natural by Taylor (1968). Sciences (ANSP) 4927, was reported as hav- ing the locality datum “Guiana” (Taylor 1968). GEOGRAPHIC DISTRIBUTION. This The single specimen known from Ecuador, aquatic species has a distribution that cov- United States National Museum (USNM) ers areas in the Amazon and Orinoco drain- 811, was collected in “Pucayacú entre Mon- age systems of South America, including the talvo et Sarayacú Río Bobonaza, Pastaza countries of Columbia, Ecuador, Peru, Ven- Province” (Taylor and Peters 1974:337). The ezuela, Brazil, and the territory of French locality information for American Museum Guiana (Alves-Silva et al. 2017; Lescure and of Natural History (AMNH) 49978 was de- Renous 1988; Maciel and Hoogmoed 2011; scribed as needing confirmation because the Oliveira et al. 2012). In Brazil, the species is specimen was reported from the Atrato River known from the states of Acre, Amapà, Am- system in Colombia (see locality 3?) (Taylor azonas, Goiàs, Maranhão, Mato Groso, Parà, 1968). A map detailing the distribution of the Rondônia, and Roraima (Caramaschi and species in Colombia was provided by Lynch Pombal 2000; Cintra et al. 2010; Oliveira et (2000). The southern distribution limit of the al. 2012). The presence of the species in Ma- species is poorly known (Oliveira et al. 2012). ranhão (Cintra et al. 2010) (locality 29 on The distribution of Potomotyphlus kaupii distribution map) seems to have been over- likely extends into northern Bolivia via con- looked in previously published distribution necting river systems. The species could also maps (e.g., Maciel and Hoogmoed 2011; possibly be found in Suriname and Guyana Oliveira et al. 2012). The type locality of the (Frost 2018). synonymized species Chthonerpeton micro- cephalum was reported as Mato Grosso (A. FOSSIL RECORD. None. Miranda-Ribeiro 1937) (see locality 15?), however, it was later noted that the specimen PERTINENT LITERATURE. Topics ad- probably was collected in an area that was dressed in the literature include: collections then the Federal Territory of Rondônia (now holdings/inventories (Bauer et al. 1993; 10

Böhme 2010; Böhme and Bischoff 1984; 2005; Hoogmoed 1979; Nussbaum and Hoog- Mendes-Pinto et al. 2011; P. de Miranda-Ri- moed 1979; Señaris and MacCulloch 2005), beiro 1955), conservation status (Coloma Peru (Catenazzi and Venegas 2012; Morales et al. 2011; ICMBio 2015; Maciel and Hoog- 1995), and Venezuela (Barrio Amorós 1998, moed 2011; Stuart et al. 2008; Wilkinson et al. 2004, 2009; Gines 1959; Gorzula and Señaris 2010), diet (Fuhrmann 1914; ICMBio 2015; 1999; La Marca 1997; Pefaur 1992; Péfaur et Presswell et al. 2002), disease (Churgin et al. al. 1992; Señaris 2004a, 2004b). Other ac- 2013; Flach et al. 2019; Hartigan et al. 2016 counts include a brief listing or mention [myxozoan infections]; Rendle et al. 2015; Ta- without a specific locality:Acosta-Galvis et mukai et al. 2014 [Batrachochytrium dendro- al. (2019), Azpelicueta et al. (1987), Bolaños batidis detection]), geometric morphomet- et al. (2008), Boulenger (1891, 1968), Budzik ric analysis of the skull (Bardua et al. 2019; and Żuwala (2011), Deban et al. (2001), Du- Sherratt et al. 2014), , capture, and bois (2005), Duellman (1981, 1993, 1999), behavior (Dunn 1942; ICMBio 2015; Kup- Duellman and Trueb (1986, 1994), Dunaev fer et al. 2006; Maciel and Hoogmoed 2011; (1999), Dünker et al. (2000), Frost (1985), Marty et al. 2007a; Rendle et al. 2015; Tay- Frost et al. (2006), Giri et al. (2004), Glaw et al. lor 1968), juvenile forms and reproduction (1998, 2000a, 2000b), Gorham (1962, 1974), (Murphy et al. 1977; Sedgwick County Zoo Gower et al. (2007), Harding (1983), Herrel 2013; Wake 1992; Wilkinson and Nussbaum and Measey (2010), Hofer (1998), Hofrichter 1998, 1999, 2006), molecular phylogenetic (1998, 2000a, 2000b), Hutchins et al. (2003), analysis and molecular data (Maciel et al. Jared et al. (1999), Jenkins et al. (2007), Lavil- 2016; San Mauro et al. 2014), morphological la and Cei (2001), Maciel et al. (2015), Maerker evolution (Wilkinson and Nussbaum 1997), et al. (2016), Matsui (1993), O’Reilly (1996), morphological comparisons with other ty- O’Reilly et al. (2002), Pinheiro (2001), Porter phlonectids (Hoogmoed et al. 2011; Maciel (1972), Pough et al. (2016), Pyron and Wiens et al. 2016; Nussbaum and Wilkinson 1989, (2011), W. Smith (1877), R. Smith and H. 1995; Wilkinson 1988, 1989, 1996; Wilkin- Smith (1973), Sokolov (1988), Teodecki et al. son et al. 1998), morphometric data on body (1998), Vitt and Caldwell (2009, 2014), War- and vertebral proportions (Renous and Gasc beck (2002), Zhao et al. (1993, 1998), and Zug 1989), and physiology (Maddin and Sherratt et al. (2001). 2014; Wells 2007; Wilkinson and Nussbaum 1997). Additional publications include a brief NOMENCLATURAL HISTORY. The spell- listing or mention for a given country: Bra- ing “Potomotyphlus kaupi,” with a single “i” zil (Atadeu Moreira 2015; Bernarde 2012; in the specific epithet, was used by Frank and Bokermann 1966; Brcko et al. 2013; de Fraga Ramus (1995). The same spelling was used by et al. 2018; de Souźa and Aveline 1994; Galat- Mitchell (2017) and Wrobel (2004). ti et al. 2007; Himstedt 1996; ICMBio 2016; Leitão 1943; Moreira 2015; Neckel-Oliveira REMARKS. The counts and measurements et al. 2012; Nussbaum and Wilkinson 1987; of the type of Caecilia dorsalis (Peters 1877) Pinheiro et al. 2012; Vaz-Silva et al. 2015; were reported to be close to those of the Vogt and Bernhard 2003), Colombia (Acos- specimen “Berlin 9092” [=ZMB 9092] by ta-Galvis 2000; Galeano et al. 2006; Hernán- Dunn (1942). Although the specimen “Berlin dez-Cuadrado et al. 2008; Lynch 2000, 2006, 10104” is labeled “type” for Caecilia dorsalis, 2007; Ruiz-Carranza et al. 1996; Señaris and both specimens should probably be consid- Acosta-Galvis 2014), Ecuador (Almendáriz ered cotypes (Dunn 1942). 1991; Coloma 1991; Miyata 1982), Guianas The misspelling of Chthonerpeton micro- (Dewynter et al. 2008; Hollowell and Reynolds cephalum by P. de Miranda-Ribeiro (1955) Account 924 11 was first noted by Taylor (1968), however, be similar to that in Typhlonectes (Tapley et the citation for P. de Miranda-Ribeiro (1955) al. 2019). There is video from the Sedgwick was listed in error by Taylor (1968) under the County Zoo (Kansas, United States) of a cap- name A. de Miranda-Ribeiro in the bibliog- tive Potomotyphlus kaupii giving birth to live raphy. young (Sedgwick County Zoo 2013). A type specimen was not specified by There is a specimen ofPotomotyphlus Taylor (1968) in his description of Potomoty- kaupii discovered during a taxon search on phlus kaupii, however he described the spe- VertNet, USNM 30534, that has a locality cies based on AMNH 42853, a male specimen datum of “” which is probably incor- collected from Iquitos, Peru. In a table of data rect considering that Belize is well out of the and measurements given by Taylor (1968: known range for the species (VertNet 2019). 262) for Potomotyphlus kaupii, one specimen was listed as “Type Rio 539.” This specimen ADDITIONAL VERNACULAR NAMES. is likely the type specimen for Chthonerpeton In English, vernacular names include Kaup’s microcephalum because they have the same caecilian (Frank and Ramus 1995, Wrobel specimen number and the same reported 2004) and (Kaup’s) Retaw (Mitchell 2017). length (A. de Miranda-Ribeiro 1955). The name “Retaw” could be a reference to the Morphological evolution between Poto- aquatic lifestyle of the species because “retaw” motyphlus and Atretochoana is relatively un- is “water” spelled backwards. Other common known (Wilkinson and Nussbaum 1997). names assigned to species that have back- Although black-and-white photographs wards spellings were used by Mitchell (2017). of the skull were published by Taylor (1969a, In French, the vernacular name appears as 1969b), the same specimen (University of Illi- Potamotyphle de Kaup (Wrobel 2004), in nois Museum of Natural History [UIM] 787) Spanish, it appears as Cecilia de Río (Wilkin- appears in both publications but has bone son et al. 2010), in Brazilian Portuguese, outlines that differ between photographs it appears as cobra cega d´água (ICMBio (Wake 2003). 2015), and in Cyrillic, it appears as речные Little information is known of the ecol- червяги = Potomotyphlus = “river worm” and ogy and physiology of Potomotyphlus kaupii, амазонская червяга = Potomotyphlus kaupii however the left may serve as a hydro- = “Amazonian worm” (Sokolov 1988). static organ and it is unlikely that this species burrows (Wilkinson and Nussbaum 1997). ETYMOLOGY. The namekaupii is in hon- Potomotyphlus kaupii has been inferred to or of Johann Jakob von Kaup (Beolens et al. be viviparous based on the reproductive mode 2013). in closely related taxa (Wilkinson and Nuss- baum 1998). Viviparity and fetuses of Poto- ACKNOWLEDGMENTS. I thank C. J. Bell motyphlus have been reported (Wake 1992), and T. J. LaDuc for encouraging me to pro- however this was questioned on the basis that duce this account. Thanks to J. Horowitz for the report did not include documented fetus- modifications to the map. My thanks also es of the species (Wilkinson and Nussbaum goes to C. Withnell and S. Scarpetta for mak- 1999). The presence of fused and sac-like fe- ing trips to photograph this species. Thanks tal gills, a morphological character that sup- to M. Wake for providing a key reference. ports the hypothesis of a monophyletic Ty- Lastly, I thank the Sedgwick County Zoo for phlonectidae, is unknown for Potomotyphlus allowing photographs to be taken of individ- (Wilkinson and Nussbaum 2006); although uals of this species in their care. the gill structure in neonates was reported to 12

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