Zootaxa 4282 (2): 395–400 http://www.mapress.com/j/zt/ Copyright © 2017 Magnolia Press
Article
ISSN 1175-5326 (print edition)
ZOOTAXA
ISSN 1175-5334 (online edition)
https://doi.org/10.11646/zootaxa.4282.2.12 http://zoobank.org/urn:lsid:zoobank.org:pub:DF66C813-9B48-4CCB-BD91-9DC6E586856B
Rediscovery and description of the holotype of Oedipus paraensis Unterstein, 1930 (Amphibia, Plethodontidae), and invalidation of the neotype ALEXANDER GUTSCHE1,2, FRANK TILLACK1 & KRISTIN MAHLOW1 1
Museum für Naturkunde, Leibniz-Institut für Evolutions- und Biodiversitätsforschung, Invalidenstr. 43, 10115 Berlin, Germany Corresponding author. E-mail:
[email protected]
2
Abstract It is proposed to invalidate the recently designated neotype for the plethodontid salamander Oedipus paraensis Unterstein, 1930, because the holotype has been rediscovered in the herpetological collection of the Museum für Naturkunde Berlin. As the taxon was recently shown being part of a species complex and the original description of O. paraensis does not provide a clear diagnosis to distinguish it from the other species, we herein supply additional morphometric data of the holotype for comparative purposes. Key words: Caudata, Bolitoglossa, neotype suppression, holotype morphology, Brazil
Introduction The tropical lungless salamander Oedipus paraensis was described by Walter Unterstein from a single specimen (Unterstein 1930). Collected by Mr. Praetorius, collector on behalf of the zoological merchant company Scholze & Pötzschke, Berlin, in "Sta. Isabel bei Para" [Santa Isabel near Pará (Belém), Brazil]. The holotype was catalogued on 17 October 1929 by Ernst Ahl—curator of amphibians and reptiles at the Zoologisches Museum Berlin (ZMB) during this time—with the registration number ZMB 32907. Taylor (1939) suppressed the generic name Oedipus Tschudi (1838) because it was preoccupied by the orthopteran genus Oedipus Berthold (1827). Taylor adopted the generic names Thorius Cope (1869) for certain small Mexican species, and Bolitoglossa Duméril, Bibron & Duméril (1854) for the remaining species formerly associated with Oedipus. In a subsequent publication, Taylor (1944) recognized O. paraensis therefore as Bolitoglossa paraensis (Unterstein, 1930). Brame & Wake (1963) synonymized B. paraensis with B. altamazonica. They argued that the holotype of O. paraensis was lost, and that Unterstein’s original description did not contain any information to distinguish paraensis from altamazonica. Furthermore, being that altamazonica was known from the vicinity of Belém, it was proposed that all extant museum specimens from that area were assignable to a single species, with B. altamazonica as the senior synonym (Brame & Wake 1963). The information "type lost" was probably provided by Heinz Wermuth (see Acknowledgments in Brame & Wake 1963), curator of herpetology at ZMB until 1961. The taxon name paraensis in combination with Oedipus or Bolitoglossa was neither mentioned in Frost’s first amphibian checklist (Frost 1985), nor in the addition subsequently published by Duellmann (1993). Bauer et al. (1993) mentioned O. paraensis in their type catalogue of caecilians and salamanders in the ZMB collection, but they also stated the holotype as "lost or destroyed". Based on a new phylogenetic analysis of the genus Bolitoglossa, Parra-Olea et al. (2004) revalidated the taxon paraensis, and removed it from synonymy with B. altamazonica. In the most recent study on Bolitoglossa from Brazilian Amazonia, Brcko et al. (2013) revised the status of B. paraensis based on extensive material collected in the Belém area. Following Brame & Wake (1963), and Bauer et al. (1993), they also considered the holotype of O. paraensis as lost, and designated a topotypical neotype (MPEG 31682).
Accepted by M. Vences: 28 Apr. 2017; published: 26 Jun. 2017
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The holotype, however, was not lost at the time when Brcko et al. (2013) prepared their study. It was rediscovered by the first author of this article in March 1998 among the uncatalogued material of the herpetological collection (Figure 1). The salamander was in the original jar with an asterisk on the jar label indicating that specimen as type. The type was then returned to the main collection, and the information was made available in the museum documentations. In addition, the existence of the type (along with two images) is also mentioned since 2004 on the online database system SysTax (Hoppe et al. 2015), the German portal of the Global Biodiversity Information Facility (GBIF). Brame & Wake (1963) and Brcko et al. (2013) pointed out that the original description of O. paraensis is too vague for a good diagnosis of the species. Therefore, we herein present new additional morphological data of the holotype of O. paraensis in comparison with data of Unterstein’s (1930) original description, following the specifications and terminology as provided by Brcko et al. (2013) for their neotype description.
Material and methods Morphological measurements were taken with a digital caliper (to the nearest 0.1 mm) and a light microscope fitted with a 10x ocular micrometer. To obtain counts of teeth by a non-invasive procedure, the specimen was subjected to micro-tomographic analysis (Phoenix nanotom X-ray|s tube; GE Company; USA) at 50 kV and 250 µA, generating 1440 projections with 750 ms per scan. Effective voxel size reached 3.9 µm. The cone beam reconstruction was performed using the datos|x-2-reconstruction software (GE Sensing & Inspection Technologies GmbH phoenix|x-ray datos|x 2.0), and the data were visualized in VG Studio Max 3.0 (Volume Graphics GmbH). The following measurements were taken according to Brcko et al. (2013): SL—standard length, snout to posterior point of vent; SVL—snout to anterior point of vent; HL—head length between tip of snout and posterior margin of mouth; HW—head width at widest point, HD—head depth at posterior angle of jaw; SGF—snout to gular fold; EYW—eyelid width; EYL—eyelid length; SNL—snout length, SP—projection snout beyond mandible; LWS—width of snout at widest point; SWS—width of snout at narrowest point; EN—distance between external nares; OD—horizontal diameter orbit; DBE—distance between anterior corners eyes; IDE—interorbital distance; WMG—width mental gland; LMG—length mental gland; NGGF—distance between nuchal groove and gular fold; SA—snout to axilla; AG—axilla to groin; PECW—pectoral width; FL—length forelimb, from axil to tip of longest finger; HLL—length hind limb, from groin to tip of longest finger; HDW—hand width; FW—foot width; LIIIF—length third finger; LIIIT—length third toe; LVT—length fifth toe; TL—tail length; TW—tail width at base; TD—tail depth at base; VL—vent length, equals SL minus SVL. Dentition: PT—premaxillary teeth; MT—maxillary teeth; VT—[anterior] vomerine teeth; DT—dentary teeth. Additional abbreviations and acronyms: PVT—posterior vomerine teeth; SNH—snout height; MPEG— Museu Paraense "Emilio Goeldi", Zoologia, Belém, Brazil; ZMB—Museum für Naturkunde [formerly Zoologisches Museum Berlin], Leibniz Institute for Evolution and Biodiversity Science, Berlin, Germany.
Results General diagnosis of holotype (Figure 1; Table 1). Presumably adult male (see discussion); specimen showing partial shrinkage due to long-term preservation; body slender, posterior part strongly curved; tail missing, only basal remnant present, tail base damaged, but basal constriction recognizable; total length 40 mm; snout rounded in dorsal aspect, sloping ventrolaterally and projecting in front of upper jaw; snout length shorter than snout width at narrowest or at widest point; snout length equal to horizontal orbit diameter; snout length shorter than distance between anterior corners of eyes but longer than snout height; nares closer to snout tip (0.2 mm) than to eyes (0.8 mm); distance between external nares less than interorbital distance; head elongate; canthus rostralis slightly rounded, but distinct; eyes and eyelids shrunken; mental gland not visible; nuchal grooves not visible; gular fold well-defined; five costal folds discernable on the right posterior trunk, additional costal folds not visible; limbs relatively slender and short; digits completely webbed, but webbing slightly shrunken between fingers and toes, fingers and toes flattened, third finger of both forelimbs and third toe of both hind limbs longest; skin smooth. For detailed measurements see Table 1.
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TABLE 1. Morphometric data (in mm) of the holotype of Oedipus paraensis Unterstein, 1930 and the neotype designated by Brcko et al. (2013). Differences in bilateral values are given in left\right order. Corrected measurements by Isabela Brcko for the neotype are given in squared brackets. See material and method section for abbreviations. Character
Holotype ZMB 32907 (own measurements)
Neotype MPEG 31682 (Brcko et al. 2013)
Remarks
SL
38.2
37.7
SVL
37.6
35.2
HL
5.3
6.0
HW
4.9
5.8
HD
3.0
3.3
SGF
8.4
2.5* [8.6]
*SGF can’t be shorter than HL
EYW
–*
1.4
*shrunken, not clear discernable
EYL
2.4\2.5
2.5
SNL
1.7
2.8
SP
0.6
1.2
LWS
3.5
3.0* [4.0]
*LWS can’t be smaller than SWS
SWS
2.8
4.8* [2.6]
*see LWS
SNH
1.1
–*
*not given
EN
1.2
1.8
OD
1.7
1.9
DBE
3.0
3.8
IDE
2.6
1.8
WMG
–*
2.7
*mental gland not discernable
LMG
–*
3.0
*see WMG
NGGF
–*
2.5
*nuchal groove not discernable
SA
11.5
12.3
AG
20.0\26.0
20.0
PECW
4.3
4.3
FL
7.4\7.3
7.1
HLL
7.5\7.6
8.1
HDW
2.3
2.3
FW
2.5
3.3
LIIIF
2.0
2.5
LIIIT
2.5
2.8
LVT
1.8
2.5
TL
1.8*
34.7
*only tail remnant exist
TW
–*
3.1
*tail base is damaged
TD
–*
3.4
*see TW
VL
0.6
2.5
PT
3
2
MT
19\20
13–14
VT
11\12
7–7
DT
27\25
33–29
PVT
118\110
–*
HOLOTYPE OF OEDIPUS PARAENSIS
*not given
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Color in ethanol. Dorsal surfaces of head and body grayish-brown to dark brown with tiny brown spots on the anterior body part; flanks anteriorly and posteriorly of the forelimbs pale brown to creamy with tiny brown spots; ventral surfaces of head and body medium to dark brown with tiny brown spots from chin to mid-body. Description of dentition (left\right order; Figure 2). Pedicels of missing teeth are included in the count. Three premaxillary teeth, enlarged, slightly bicuspid, conical; 19\20 maxillary teeth, bicuspid, monostichous, conical, mostly shorter than premaxillary teeth, but longest reach similar size as premaxillary teeth, anterior 10%\12% and posterior 25%\23% of maxillary bone without teeth; 11\12 anterior vomerine teeth bicuspid and of similar size, polystichous, conical, longest of similar size as premaxillary teeth, lateral 39%\31% of the vomer without teeth; 118\110 bicuspid posterior vomerine teeth, associated with the parasphenoid, slightly obliquely directed backwards, conical, gradually increasing in length medially and posteriorly, longest posteriorly, longer than premaxillary teeth, closely grouped in two patches as acute-angled triangles, not touching the anterior vomerine teeth; 27\25 bicuspid dentary teeth, monostichous, conical, longest of similar size as premaxillary teeth, gradually decreasing in size posteriorly, anterior 4%\3% and posterior 39%\40% of dentary bone without teeth.
FIGURE 1. Holotype of Oedipus paraensis Unterstein, 1930 (ZMB 32907); scale bar 5 mm.
Discussion The recent condition and measurements of ZMB 32907 match quite well with the original description given by Unterstein (1930) confirming its identity as the holotype of O. paraensis. Some data are equal to Unterstein, e.g., missing tail, total length, SGF ["4½ mal in der Länge von der Schnauzenspitze bis zum After"; 4½ times of SVL], or SNL = OD (see Table 1). Probably due to shrinkage in ethanol and/or partly drying-out, some characters differ slightly, such as HW ["Kopfbreite 6¼–6¾ mal in der Länge von der Schnauzenspitze bis zum After"; head width 6¼–6¾ times of SVL], which is now 5 mm (at widest point) vs 5.6–6 mm, and some characters are not visible anymore like the nuchal groove or the majority of the costal folds. In contrast to Unterstein (1930), the posterior vomerine teeth (or vomerine tooth patches) are not touching the anterior vomerine teeth ["Parasphenoidzähne in einer großen Gruppe an die Vomerzähne stoßend"] (Figure 2). Unterstein (1930) did not mention the sex of the holotype. Characters for sex determination like the mental gland are not visible, and body dimensions lie within the ranges of both sexes as given by Brcko et al. (2013). However, we assume the holotype to be a male, based on some dentition characters that commonly occur in male plethodontid salamanders (e.g., Ehmcke & Clemen 2003, Ehmcke et al. 2004). In contrast to females, males often have enlarged teeth in the front of the maxilla, and their premaxilla is not aligned with the maxilla showing a gap on both sides.
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Despite some shrinkage, the measurements of the holotype largely correspond with the data of the neotype (Table 1), and lie within the ranges of the data sets given by Brcko et al. (2013: Table 3) for the whole sample of B. paraensis. Some discrepancies occur in the paper of Brcko et al. (2013): The measurements of the forelimb length (FL) and hind limb length (HLL) are explained in the method section as being taken from the axil and groin respectively to longest fingertip and toetip, but this does not agree with their drawing in figure 3, where FL is shown as taken from the shoulder and HLL from the base of the tail. However, we took the measurements as explained in the text. The distance from the snout to the gular fold (SGF) of the neotype is given with a length of 2.5 mm and therefore is considerably shorter than the head length (HL), which can’t be possible (Table 1). Snout width at the widest point (LWS) of the neotype is mentioned as being shorter (3.0 mm) than at the narrowest point (SWS; 4.8 mm), which again is not possible; additionally, the latter value is even higher than the maximum value (3.8 mm) given for the whole sample of B. paraensis by Brcko et al. (2013). To clarify these discrepancies in Brcko et al. (2013), we contacted the authors, and they agreed that there were some mistakes and made some new measurements that are herein provided in Table 1. The holotype of O. paraensis is available in the ZMB collection in a condition, which enabled us to provide additional morphometric data for comparative purposes. According to Article 75.8 of the International Commission on Zoological Nomenclature (ICZN 1999), the rediscovered holotype again becomes the namebearing type and the neotype is set aside in our case. During World War II, the ZMB amphibian and reptile collection was sheltered in the basement of the building. It is most likely that the type of O. paraensis somehow ended up among the uncatalogued material during that time. However, since the post-war period, the herpetological collection has been reorganized several times and numerous types stated as "lost", "not traced" or "presumably in ZMB" in literature (e.g., Frost 1985, 2016) have been recovered. It is therefore recommended that anyone interested in the current status of any ZMB material contact the collection staff directly to enquire about any particular specimens, or look at the SysTax site (Hoppe et al. 2015).
FIGURE 2. Skull of the holotype of Oedipus paraensis (ZMB 32907), CT scan. A: cranium ventral view, mandible and vertebral column virtually removed; B: mandible dorsal view, cranium virtually removed; scale bar 1 mm.
Acknowledgments We are much obligated to Marinus Steven Hoogmoed (MPEG, Brazil) for his valuable comments to an earlier version of the manuscript, and to Isabela Brcko (Belém, Brazil) for preparing corrected measurements of the neotype. We also thank James R. McCranie (Miami, USA) for some details on the dentition of Bolitoglossa, and the two reviewers for their comments that improved the quality of the text.
HOLOTYPE OF OEDIPUS PARAENSIS
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Literature cited Bauer, A.M., Good, D.A. & Günther, R. (1993) An annotated type catalogue of the caecilians and salamanders (Amphibia: Gymnophiona and Caudata) in the Zoological Museum, Berlin. Mitteilungen aus dem Zoologischen Museum in Berlin, 69, 285–306. https://doi.org/10.1002/mmnz.19930690208 Berthold, A.A. (1827) Latreille’s natürliche Familien des Thierreichs. Aus dem Französischen. Mit Anmerkungen und Zusätzen von Dr. Arnold Adolph Berthold. Im Verlage des Gr. H. S. priv. Landes-Industrie-Comptoirs, Weimar, x + 606 pp. Brame, A.H. Jr. & Wake D.B. (1963) The salamanders of South America. Contributions in Science. Los Angeles County Museum, 69, 1–72. Brcko, I., Hoogmoed, M.S. & Neckel-Oliveira, S. (2013) Taxonomy and distribution of the salamander genus Bolitoglossa Duméril, Bibron & Duméril, 1854 (Amphibia, Caudata, Plethodontidae) in Brazilian Amazonia. Zootaxa, 3686 (4), 401– 431. https://doi.org/10.11646/zootaxa.3686.4.1 Cope, E.D. (1869) A review of the species of the Plethodontidae and Desmognathidae. Proceedings of the Academy of Natural Sciences of Philadelphia, 1869, 93–118. Duellmann, W.E. (1993) Amphibian Species of the World: Additions and Corrections. The University of Kansas, Museum of Natural History, Special Publication, 21, iii + 372 pp. Duméril, A.M.C., Bibron, G. & Duméril, A.H.A. (1854) Erpétologie Genérale ou Histoire Naturelle Complète des Reptiles. Tome Neuvième. Librairie Enclyclopédique de Roret, Paris, iv + 440 pp. Ehmcke, J. & Clemen, G. (2003) The skull structures of six species of Mesoamerican plethodontid salamanders (Amphibia, Urodela). Annals of Anatomy, 185, 253–261. https://doi.org/10.1016/S0940-9602(03)80035-3 Ehmcke, J., Wistuba, J. & Clemen, G. (2004) Gender-dependent dimorphic teeth in four species of Mesoamerican plethodontid salamanders (Urodela, Amphibia). Annals of Anatomy, 186, 223–230. https://doi.org/10.1016/S0940-9602(04)80007-4 Frost, D.R. (1985) Amphibian species of the world. A taxonomic and geographical reference. Allen Press inc. and ASC, Lawrence, Kansas, 732 pp. Frost, D.R. (2016) Amphibian Species of the World: an Online Reference. Version 6.0. Available from: http:// research.amnh.org/herpetology/amphibia/index.html (accessed 24 February 2017) Hoppe, J.R., Boos, E., Fischer, S., Vogelpohl, C. & Stützel, T. (2015) SysTax - a Database System for Systematics and Taxonomy. Available from: http://www.biologie.uni-ulm.de/cgi-bin/query_all/query_all.pl?query=oedipus+paraensis (accessed 24 February 2017) ICZN (1999) International Code of Zoological Nomenclature. 4th Edition. The International Trust for Zoological Nomenclature, London, 306 pp. Parra-Olea, G., García-París, M. & Wake, D.B. (2004) Molecular diversification of salamanders of the tropical American genus Bolitoglossa (Caudata: Plethodontidae) and its evolutionary and biogeographical implications. Biological Journal of the Linnean Society, 81, 325–346. https://doi.org/10.1111/j.1095-8312.2003.00303.x Taylor, E.H. (1939) New Salamanders from Mexico with a discussion of certain new forms. The University of Kansas Science Bulletin, 26, 407–439. Taylor, E.H. (1944) The genera of plethodont salamanders in Mexico, Pt. I. The University of Kansas Science Bulletin, 30, 189– 232. https://doi.org/10.5962/bhl.part.6508 Tschudi, J.J. (1838) Classification der Batrachier mit Berücksichtigung der fossilen Thiere dieser Abtheilung der Reptilien. Buchhandlung von Petitpierre, Neuchâtel, 99 + i pp., pls. 1–6. https://doi.org/10.5962/bhl.title.4883 Unterstein, W. (1930) Beschreibung zweier neuer Molche der Gattung Oedipus. Zoologischer Anzeiger, 8, 270–272.
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