Herpetology Notes, volume 10: 89-99 (2017) (published online on 01 March 2017)

Identity and history of an odd, old gecko (Gymnodactylus geckoides Spix, 1825, Phyllodactylidae) from the Zoologische Staatssammlung München, Germany

Herbert Rösler1, Frank Glaw2 and Hinrich Kaiser3,*

Abstract. We describe a gecko specimen that was initially catalogued in the Zoologische Staatssammlung München, Germany, under the name Gymnodactylus marmoratus, a species now known as Cyrtodactylus marmoratus from Java, Indonesia. A close inspection revealed that this very old specimen (catalogue number ZSM 245/0) is actually a member of the South American species Gymnodactylus geckoides. Intensive research of historic documents initially appeared to indicate that the specimen could be a lost Spix syntype, yet ultimately we were unable to pinpoint the specimen’s origin. Nevertheless, its provenance in the early part of the 19th Century makes for an intriguing historical herpetological anecdote that once again underlines the importance of active research in historical natural history collections.

Keywords. Phyllodactylidae, Gymnodactylus geckoides, Spix, Leuchtenberg, Brazil, gecko.

Introduction use for this taxon. The species identification on the jar label differs by the species authority, which is listed The herpetological catalogue of the Zoologische using another author, as “Gymnodactylus marmoratus Staatssammlung München, Germany (ZSM) shows (Fitz.)” (Fig. 3). The abbreviation “Fitz.” references under the accession number ZSM 245/0 a gecko listed the Austrian herpetologist Leopold Josef Fitzinger as “Gymnodact. marmoratus (Kuhl)” with locality (1802–1884). In the following paragraphs, we describe “Java” (Fig. 1). The corresponding original entry into this rather mysterious specimen, check the initial the collection’s permanent card index catalogue, in identification, and report on our attempts to trace its the same handwriting, was initially identical to that in origin and its history. the catalogue (Fig. 2). Subsequently, a correction was made to the species authority in the card entry, changing Materials and Methods “(Kuhl)” to “(Gray, 1831)” (Fig. 2), the correct authority for Cyrtodactylus marmoratus and the name in current We collected a variety of morphological data to ascertain the identity of ZSM 245/0 (Fig. 4), an adult male gecko specimen in poor condition. In order to compare the characteristics of this specimen with other geckos, we completed a series of measurements and compiled 1 Herbert Rösler, Senckenberg Naturhistorische Sammlungen several scale counts. Measurements were taken to the Dresden, Museum für Tierkunde, Königsbrücker Landstraße 159, 01109 Dresden, Germany. nearest 0.1 mm using digital calipers. Scale counts were E-mail: [email protected] completed using a dissecting stereomicroscope. All data 2 Frank Glaw, Zoologische Staatssammlung München (ZSM- were taken on the right side of the specimen, unless SNSB), Münchhausenstraße 21, 81247 München, Germany. stated otherwise. We report the scale count on both E-mail: [email protected] sides of the specimen by listing scales as R/L, where 3 Hinrich Kaiser, Department of Biology, Victor Valley College, R denotes the right and L the left side. The number of 18422 Bear Valley Road, Victorville, California 92395, USA; subdigital lamellae was counted on fingers (F) and toes and Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution. Washington, D.C. (T) in the usual a basal-to-apical direction on both right 20013, USA. and left digits, with the number of enlarged subdigital * Corresponding author e-mail: [email protected] lamellae given in parentheses. Thus, a lamellar count of 90 Herbert Rösler et al.

Figure 1. Original entry for ZSM 245/0 as “Gymnodact. marmoratus (Kuhl)” in the catalogue of the Zoologische Staatssammlung München, Germany.

12 with four enlarged lamellae on the first finger of the Description of ZSM 245/0 right hand, with the first finger on the left hand missing, Sex.—Adult male. is reported as F1 = 12(4)/–. In some cases scales could not be counted reliably because of the condition of the Measurements (in mm).—Snout–vent length 43.5, tail specimen, and such values are preceded by “ca.” length 56.0, head length 10.5, head width 8.1, head depth 6.1, distance from the tip of the snout to the anterior edge of the orbit 4.6, distance from the posterior edge Results of the orbit to the posterior edge of the ear opening 4.4, Below, we provide relevant morphological orbital diameter 2.1, diameter ear opening 1.2, rostral characteristics of ZSM 245/0 to facilitate the compari- width 2.1, rostral height 0.9, mental width 3.3, mental sons subsequently presented. length 2.4. Identity and history of an odd, old gecko from the ZSM, Germany 91

Figure 2. Card index catalogue record of Gymnodactylus marmoratus in the Zoologische Staatssammlung München, Germany. Based on the order of entries (beginning at the top of the card), ZSM 245/0 must have been added in 1909 or thereafter. Note the corrections of species author from Kuhl to Gray 1831, and the reference at the top “siehe jetzt Cyrtodactylus” [see now Cyrtodactylus] to flag the change in genus. Red check marks next to specimen numbers indicate that these specimens were present in the collection during an inventory to determine which specimens survived the Second World War.

Figure 3. Label from the jar, in which specimen ZSM 245/0 is housed. The correction indicates that this type of label was initially produced to identify the collection of Lorenz Müller. In this case, one of Müller’s labels was altered to indicate its use for a specimen of the “old collection,” which pre-dated Müller. The taxon authority, given as “Fitz” to indicate Leopold Josef Fitzinger, is in error.

External characters and pholidosis.—Head slightly infralabials, 3/3 nasals, one internasal. Nares in contact flattened, distinct from body. Pupil round with smooth with rostral and first supralabial, nasorostrals twice as edge. Rostral with a short, incomplete suture separating large as postnasals. Snout not medially concave, snout the upper thirds of the scale, 7/6 supralabials, 5/5 scales medially twice as large as occipital granular 92 Herbert Rösler et al.

Figure 4. ZSM 245/0, originally catalogued as Cyrtodactylus marmoratus (Gray, 1831) from Java, and now reidentified as Gymnodactylus geckoides Spix, 1825 from Brazil.

scales. Canthal and interorbital scales of the same smooth, imbricate. Granular scales on the posterior size. 13/14 scales between postnasals and orbit, ca. 33 surface of the forelimbs as large as occipital scales. scales between fourth supralabials, ca. 24 interorbitals. Scales on the upper leg anteriorly and dorsally as large Occipital scales granular, no tubercles in the interorbital as ventrals, flat, smooth, imbricate. Granular scales on and occipital area. Nuchals granular, nuchal and the posterior surface of the hind limbs as large as the temporal tubercles 1.5–2 times as large as bordering medial snout scales. Tubercles absent on fore- and hind granular scales, round, conical, blunt to pointed. Mental limbs. Toes not expanded, bent. Number of subdigital U-shaped, wider than rostral. Two postmentals, as lamellae F1 = 12(4)/–, F4 = 13(5)/–, T1 = 9(2)/9(2), large in area as dorsal tubercles, triangular, medially T4 = 17(8)/17(8). T1–T4 connected basally by narrow separated by eight granular gular scales, in contact with webbing. 1st infralabials; no well-defined, differentiated lateral Tail round in cross-section, with several indistinct postmentals; scales bordering infralabials 3–4 times as tubercular whorls occurring on its proximal third. 1/1 large as occipital scales; throat scales granular, as large cloacal tubercles. Tail scales dorsally twice as large as as occipital granules. dorsal scales, laterally three times as large, flat, keeled, Dorsal and lateral scales as large as occipital scales, imbricate, with a pointed posterior edge, in regular domed, smooth, non-overlapping. Dorsal and lateral whorls across the tail. Whorls 1–7 ventromedially tubercles in 13 regular longitudinal rows; tubercles separated by seven scales, tail tubercles in the first triangular, conical, with a simple keel, bordered by 8– whorl 1.5 times the size as dorsal tubercles, triangular, 11 granular scales, with no (rarely one) granular scales longer than wide, weakly conical, keeled, pointed, size separating tubercles longitudinally and 1–3 granular and number of tubercles declining continuously towards scales separating tubercles on their sides. Dorsal the tail tip. Whorl 1 with six, Whorl 2 with five, and tubercles extending close to the vertebral line, with 5–7 Whorls 3–8 with two tubercles, no tubercles present granular scales separating the two medial tubercle rows in the more distal parts of the tail. Medial subcaudals across the vertebral line; ca. 47 paravertebral tubercles. enlarged, flat, smooth, imbricate, running along the tail in a single longitudinal row. Lateral fold absent. Belly scales twice as large as dorsal tubercles, flat, smooth, imbricate; 14 ventrals across Colouration and pattern.—Dorsally and ventrally the venter at mid-body. Pre-cloacal and femoral pores light olive green, patterning sepia. Dorsal flecks small, absent. indistinct, in longitudinal rows. Upper part of tail with Scales on the upper and lower arm both anteriorly distinctly separated, narrow, dark rings, in the distal and posteriorly as large as the scales on the chest, flat, third with five light rings. Identity and history of an odd, old gecko from the ZSM, Germany 93

Remarks.—The specimen is damaged. The injuries Comparisons with the genus Cnemaspis Strauch, include missing or loose skin both dorsally and ventrally, 1887.—Round pupils, as evident in ZSM 245/0, exist which appears to be a result of a poorly executed among Asian geckos only in the genus Cnemaspis capture. The missing digits on the left hand, in contrast, sensu lato (Smith, 1933; Underwood, 1954; Gamble are clearly the result of an earlier injury. The edges of et al., 2015). According to Grismer et al. (2014), the those wounds are healed and covered with scales. southernmost locality for a species from the Indochina/ Sunda clade of Cnemaspis in Asia is Pulau Karimata, Comparison of ZSM 245/0 to Other Bent-Toed where a population of C. kendallii (Gray, 1845) occurs. Geckos The species C. timoriensis (Duméril and Bibron, 1836), erroneously attributed to Timor Island, is not a member Upon discovery of ZSM 245/0 we had immediate and of the genus Cnemaspis (Rösler, 2016). significant doubts regarding its proper identification. A Species of Cnemaspis either lack dorsal tubercles multitude of morphological characteristics (including entirely, or they possess only a few spinous, spaced- size, pupil shape, dorsal tubercle arrangement, number out tubercles arranged dorsally and laterally, as well of mid-body ventral scales, toe shape) did not fit those of as a greater number of relatively small ventral scales Cyrtodactylus marmoratus. We began our comparisons (Loveridge, 1947; Perret, 1963; Spawls et al., 2002; by checking the specimen against species possessing Das, 2005; Bauer et al., 2006; Manamendra-Arachchi et similar toe morphologies known to occur on Java and on al., 2007; Trape et al., 2012; Grismer et al., 2014, 2015). other islands in the region. When this investigation did ZSM 245/0 differs from all Asian forms, as well as not produce a match with any Asian gecko species, we from the African species that at this point remain in the decided to expand our comparisons to include geckos genus Cnemaspis (see Gamble et al., 2011, 2012; Pyron without any geographic bias. et al., 2013; and Grismer et al., 2014 for additional Comparisons with Cyrtodactylus.—Four species comments on the taxonomy of the genus Cnemaspis), of Cyrtodactylus are currently known from Java, by the presence of many tubercle rows that closely align the purported collection locality of ZSM 245/0: to the vertebral region, more dorsal tubercles that are Cyrtodactylus klakahensis Hartmann et al., 2016; C. longitudinally in contact, triangular, and keeled, and marmoratus Gray, 1831; C. petani Riyanto et al., 2015, fewer, proportionally larger ventral scales. Since the and C. semiadii Riyanto et al., 2014. ZSM 245/0 can be listed characteristics preclude that this specimen could differentiated from these four species by the following be a member of the genus Cnemaspis, the round pupils characteristics (those of ZSM 245/0 are provided in of ZSM 245/0 must therefore not be a species-level parentheses): postmentals in contact (postmentals characteristic but, as sometimes occurs in other gecko separated by granular scales), 30–40 ventral scales at species with slit-like pupils, an artifact of preservation mid-body (14), subcaudals not enlarged (enlarged). In (see Cassimiro and Rodrigues, 2009: Fig. 2B). addition, ZSM 245/0 differs from C. klakahensis, C. Comparisons with Nactus.—The genus Nactus Kluge, marmoratus, and C. petani by the absence of tubercles 1983 includes several species that are morphologically on the limbs and by the absence of precloacal and very similar to ZSM 245/0. The most significant femoral pores (Rösler et al., 2007; Riyanto et al., 2015; areas for the diversity of Nactus include many islands Rösler, 2016). Cyrtodactylus semiadii has 36 ventral southeast of Wallacea (Bauer, 1994; Kraus, 2005) as scales at mid-body (14), 47 paravertebral tubercles well as Australia (Heinicke et al., 2010; Cogger, 2014). (37–40), tubercles on its lateral fold (no lateral fold, Its distribution also extends across the entire Indian no tubercles), 14–15 subdigital lamellae on the 4th toe Ocean into the Mascarene Islands (Loveridge, 1951; (17), and subcaudals that are not enlarged (enlarged) Vinson and Vinson, 1969; Arnold and Bour, 2008), (Riyanto et al., 2014). Furthermore, despite the and it is therefore not inconceivable that Nactus would relatively significant damage to the skin we were able to occur on Java. The locality of a Pacific Nactus species determine that ZSM 245/0 cannot belong to the genus closest to Java is Pulau Salibabu, North Sulawesi, Cyrtodactylus because of (1) the position, orientation, Indonesia (Koch et al., 2009). Comparison of body size and type of dorsal tubercles, which occur in regular as well as the numbers and shapes of dorsal tubercles, longitudinal rows and are usually in contact, (2) the very ventral scales, subdigital lamellae, and subcaudal scales large ventral scales, and (3) the absence of precloacal readily allows morphological differentiation of ZSM and femoral pores (see Rösler and Glaw, 2008). 245/0 from all described (as well as some still unnamed) 94 Herbert Rösler et al.

Figure 5. The original illustration of Gymnodactylus geckoides (Spix 1825: Tafel 18, Fig. 1), which Vanzolini (2004) considered the holotype (as an “iconotype”).

Nactus species (see Arnold and Jones, 1994; Zug and genus Gymnodactylus, which is currently typified only Moon, 1995; Probst, 1996; Zug, 1998; Kraus, 2005; by the “iconotype” of the original illustration (Spix, Rösler et al., 2005; Arnold and Bour, 2008; Zug and 1825: Pl. 18, 1; Fig. 5 herein). Fisher, 2012). On the Origin of ZSM 245/0 Comparisons with Gymnodactylus.—Similar attributes to ZSM 245/0 in terms of size and pholidosis During our investigation of the specimen’s origin, we exist in species of the New World genus Gymnodactylus initially considered that it could be the last remaining Spix, 1825. Comparisons showed that several species syntype of Gymnodactylus geckoides from the of Gymnodactylus possess chin shields and dorsal “zoological-zootomic collection” of Johann Baptist scales that conform to those of ZSM 245/0 (Gray, Ritter von Spix (1781–1826). We made this connection 1845; Barbour, 1925; Amaral, 1932; Vanzolini, 1982; for a variety of reasons, which derived from the Cassimiro and Rodrigues, 2009). The specimen in manner in which the ZSM acquired some of its oldest question can be distinguished from G. amarali Barbour, specimens. For example, in his list of animals collected 1925, G. guttulatus Vanzolini, 1982, and G. vanzolinii in Brazil, von Martius (1831) included 130 species of Cassimiro and Rodrigues, 2009 by dorsal tubercles amphibians (inclusive of reptiles, which were at that that are longitudinally aligned next to one another time still integrated into that class of animals), but vs. separated by small scales. It can be distinguished unfortunately he did not provide specific information from G. darwinii (Gray, 1845) by the smaller number about the specimens themselves. There is nothing of paravertebral tubercles (Cassimiro and Rodrigues, known about the number and later disposition of the 2009; da Silva Junior, 2010; Vanzolini, 1982, 2004, G. geckoides specimens collected by Spix (Hoogmoed 2005). However, the specimen agrees completely with and Gruber, 1983; Franzen and Glaw, 2007). The last G. geckoides Spix, 1825 based on the descriptions of author who directly dealt with some of the material Spix Spix (1825), Boulenger (1885), Steindachner (1867), brought to Munich was Wilhelm Carl Hartwig Peters Strauch (1887), Grantsau (2003), Vanzolini (1953, (1815–1883). Alas, the Spix collection was no longer 2004), and da Silva Junior (2010). complete even in Peters’s days (Peters, 1877). A recent Our evaluation shows that both the species designation audit of specimens in the Museum für Naturkunde, and the locality that had been assigned to this gecko Berlin, Germany (ZMB) at our request revealed that, to date are false and in need of correction. We hereby despite historic loans between Munich and Berlin, at correct the species designation and place ZSM 245/0 this time there is no specimen of G. geckoides in the into the neotropical species G. geckoides Spix, 1825. ZMB collection that could have been collected by Spix Gymnodactylus geckoides is the type species of the (R. Günther and F. Tillack, pers. comm.). It therefore Identity and history of an odd, old gecko from the ZSM, Germany 95 appears that the material loaned to Peters from Munich for how this specimen could have become part of by Carl von Siebold (1804–1885) was completely the ZSM collection. After Spix died, the holdings of returned. Additionally, an inspection of material in the South American geckos at the ZSM were expanded by Naturalis Biodiversity Center, Leiden, The Netherlands several collectors (see Gruber, 1992; Franzen and Glaw, (formerly the Rijksmuseum van Natuurlijke Historie, 2016). Substantial contributions at the beginning of the RMNH) ascertained that, beyond the material listed by 20th century were made especially by Lorenz Müller. Hoogmoed and Gruber (1983), there are no additional However, Müller started his expedition to Brazil only specimens collected by Spix in that collection (E. in 1909 (Müller, 1914) and all of his specimens from Dondorp, pers. comm.). Furthermore, the Spix collection South America were registered in the new catalogue in Munich suffered additional losses due to poor established by him in 1907. It is therefore improbable that collection management practices and the circumstance he could have contributed ZSM 245/0 to the collection. that some of the specimens were made available for On the other hand, ZSM 245/0 could have originated university teaching and may have been dissected and with the collection of Duke August von Leuchtenberg, destroyed (Hoogmoed and Gruber, 1983; Schönitzer Prince of Portugal (also known as Auguste Charles and Franzen, 2016). Thus, there did not appear to be any Eugène Napoléon de Beauharnais, 1811–1835). He simple link from this specimen to Spix. was in friendly contact with the ZSM zoologist Johann An important indicator that initially led us to identify Georg Wagler (1800–1832) until the latter’s death, ZSM 245/0 as one of the specimens collected by Spix and it is possible that Leuchtenberg received several was the fact that it is part of the “alte Sammlung” [old specimens from Spix or Wagler for his own collection. collection], which can be recognized by a running Leuchtenberg’s collection was acquired by the ZSM number, followed by a “/0”. The old collection mostly in 1858 and contained a few herpetological specimens comprises the Spix collection, but also other material without locality data from Brazil (e.g., Amphisbaena collected until 1907 (Glaw and Franzen, 2006). microcephalum, ZSM 2623/0, a species collected by According to this numbering system, established by Spix and described by Wagler) but it remains unclear then-curator Lorenz Müller (1868–1953), Catalogues 1– whether these specimens were collected by Spix or 3 of the ZSM include a majority of the amphibians and another collector. reptiles assembled by Spix, and ZSM 245/0 is included in Ultimately, our findings did not provide incontrover- Catalogue 1 (numbers 1/0 to 1824/0; Glaw and Franzen, tible evidence for a connection with Spix. We did, 2006). While it is not known why this specimen was not however, make the following determinations. part of a series of G. geckoides when it was accessioned, • Based on the early entry in Catalogue 1, it is clear it is most likely that it had become separated from one that ZSM 245/0 was collected in the 19th Century of the Brazil collections before the establishment of (Schönitzer and Franzen, 2016). The so-called old the catalogues and therefore subsequently needed to be collection includes all of the type material deposited listed as an individual. It is also possible that the reason by Spix and Wagler, except those specimens, which for its putative separation from a larger lot may have were lost or exchanged. been the poor condition of the specimen itself, or that the • ZSM 245/0 was presumably labeled as Gymnodactylus doubts surrounding the South American origin of this marmoratus due to having been received with the specimen initially somehow led to specimen separation. locality “Java.” At the time of its accession into By the same token, the distribution of a specimen series the Munich collection G. marmoratus was the only into several specimen jars was not necessarily unusual species of Gymnodactylus known from Java, even (e.g., Müller and Brongersma, 1933). Some individual though clearly at least the species G. geckoides would exemplars may also have been donated to other already have been known to Spix, the taxon’s author. collections or exchanged. This is the reason why there Lorenz Müller, who was active in the herpetological are several Spix type specimens in the RMNH (Müller division of the Zoologische Staatsammlung München and Brongersma, 1933; Hoogmoed and Gruber, 1983; beginning in 1903, appears to have catalogued newly Hoogmoed, 1986). purchased or donated specimens in his card index While the circumstances of the catalogue and catalogue before adding the old collection (see the card catalogue entries seemed to provide valuable order of entries in Fig. 2). circumstantial evidence for the age and disposition of • Many of the Brazilian Spix types are still extant in ZSM 245/0, we soon realized through our consultation the Zoologische Staatssammlung München (Fittkau, of historic documents that other possibilities existed 1992; Hoogmoed and Gruber, 1983; Gruber, 1992), 96 Herbert Rösler et al.

whereas the type series of Gymnodactylus geckoides Discussion is considered lost (Hoogmoed and Gruber, 1983; Our research into the history and provenance of a Franzen and Glaw, 2007). single specimen of uncertain identity and provenance in • Some specimens collected by Spix were donated the ZSM collection shows once again that taxonomic or exchanged to other museums, and some may treasure is almost certainly still hiding out in many simply have been used up during dissections in natural history collections. Due to the complex and anatomy classes. Some types of various species were fragmented political history of the German-speaking exchanged with the Rjiksmuseum van Natuurlijke parts of Europe, this may be particularly true for German History (now Naturalis Biodiversity Center), Leiden, natural history collections (e.g., Böhme, 2014; Mecke The Netherlands (see Müller and Brongersma, 1933; et al., 2016; Rösler and Kaiser, 2016; see Kaiser et al., Hoogmoed, 1986; Hoogmoed and Gruber, 1983). 2017), but significant findings have also been reported Among these are no specimens of G. geckoides collected by Spix (E. Dondorp, pers. comm.). recently for important collections in Italy, Sweden, the • Fitzinger (1827) was apparently able to examine Netherlands, the United Kingdom, and the United States the complete Spix collection, whereas Peters (1877) (e.g., Kathriner et al., 2014; Zug and Kaiser, 2014; noted some gaps when he borrowed the specimens Hartmann et al., 2016; Kieckbusch et al., 2016; O’Shea from Munich for his examination (a specimen of and Kaiser, 2016; O’Shea et al., 2015, 2016). Therefore, Gecko cruciger Spix, 1825 = Hemidactylus mabouia not only must we recognize the archival and historical Moreau de Jonnès, 1818 was missing). In the nature of natural history collections, but also their active collection at the Museum für Naturkunde, Berlin, contributions to our studies of biodiversity. At a time Germany, there are currently no specimens of G. when funding for fieldwork and molecular studies is geckoides that could have been collected by Spix (R. awarded via highly competitive processes, it perhaps Günther and F. Tillack, pers. comm.). behooves both researchers and funding agencies to • It is possible that ZSM 245/0 is a specimen from remember that natural history collections already hold the Leuchtenberg collection. The specimen may material for a plethora of research questions. have been collected by the royal person himself during his five-month stay in Brazil (Voigt, 1835). Acknowledgments. We thank Rainer Günther and Frank Tillack Alternatively, the specimen could have been obtained (Museum für Naturkunde, Berlin, Germany) for conducting a by Leuchtenberg’s collector father from an animal comprehensive search of their collection for Spix specimens dealer, or he may have obtained it via diplomatic and for the loan of material. We greatly appreciate the help of channels from Brazil. Esther Dondorp (Naturalis Biodiversity Center, Leiden, The • The Leuchtenberg collection includes specimens Netherlands), who confirmed that only Spix material already from several regions with a global scope. Aside reported on in publications currently exists in the Leiden from Brazilian specimens, it does include some from collection. For his work on the images of the gecko from Munich, we thank Gerd Trautmann (Laboe, Germany). For their excellent Java. For individual specimens in this collection, reviews and comments on an earlier version of the manuscript, we uncertainties regarding their correct place of origin thank Aaron Bauer, Wolfgang Böhme, and Marinus Hoogmoed. surfaced very early, which led to errors and confusion. For example, the type locality of Cercosaura ocellata Wagler, 1830 is given as “Asia?” in the original References

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Accepted by Alessandro Morais