Zoologische Mededelingen, 82 (December 2008)
A replacement name for Sphenomorphus keiensis (Kopstein, 1926) from the southeastern Moluccas, Indonesia (Reptilia: Squamata: Scincidae) with a redescription of the species
G.M. Shea 1, J.P. Michels 2
Keywords: Scincidae; Sphenomorphus; systematics; nomenclature; Indonesia.
Lygosoma keiensis Kopstein, 1926 is demonstrated to be a junior primary homonym of Lygosoma cyanogaster keiensis Sternfeld, 1918. The junior homonym, now placed in Sphenomorphus, is given the replacement name S. capitolythos, and redescribed from the holotype, the only known specimen.
Many homonyms exist among nominal species-group taxa described in the scincid genus Lygosoma, the genus to which most skinks were referred during the second half of the 19th and first half of the 20th century (Duméril & Duméril, 1851; Boulenger, 1887; De Rooij, 1915). This is particularly the case among species from southeast Asia and the Australopapuan region. Many of these homonyms are secondary, arising from Boulenger’s (1887) combination in Lygosoma of species originally described in a number of genera, and a variety of replacement species names were created at the time (Boulenger, 1887; Ahl, 1925). However, some are primary homonyms, reflecting the long period after Boulenger’s monograph when most new skink species were referred to Lygosoma.
Lygosoma was subsequently split into numerous genera, most notably by Smith (1937), Mittleman (1952) and Greer (1974). For example, the 112 species referred to Lygosoma in the most recent monographic treatment of the skinks of the Indonesian/Papuan archipelago (De Rooij, 1915) are currently assigned to 16-17 genera (there is recent disagreement about the validity of the genus Apterygodon; Mausfeld & Schmitz, 2003).
The dismemberment of Lygosoma, and the lack of any complete checklist of names in the family since Boulenger (1887), has resulted in little attention being paid in recent years to the existence of primary homonyms among species originally described in Lygosoma but now in different genera. Most of these homonyms apply to names now considered synonymous with other taxa and not in present use. Therefore there is no need to provide them with a substitute name (Article 23.3.5; International Commission on Zoological Nomenclature, 1999). However, there still remain some problematic names.
One publication particularly rife with homonyms is Sternfeld (1918), often incorrectly cited as published in 1920 (e.g., Loveridge, 1948; Tyler, 1968; Zweifel, 1979; Mys, 1988; Brown, 1991). Sternfeld described 16 new species and subspecies of skinks in this paper. Eight of these are homonyms. Sternfeld twice proposed the same species or subspecies name for two taxa in the same paper: Lygosoma (Hinulia) maindroni wolfi (p. 395) and Lygosoma (Otosaurus) wolfi (p. 397), and Lygosoma (Riopa) albofasciolatum boettgeri (p. 418) and Lygosoma (Emoa) boettgeri (p. 406), possibly considering that either the differing subgeneric assignment, or the allocation of names to subspecies rather than species, avoided the homonymy. These primary homonyms were identified by Mertens (1924), who proposed the nomina nova Otosaurus sternfeldi (currently Sphenomorphus concinnatus (Boulenger, 1887)) for L. wolfi, and Riopa albofasciolatum poehli (currently Eugongylus albofasciolatus (Günther, 1872)) for L. a. boettgeri.
Three of Sternfeld’s skinks, Lygosoma (Keneuxia) smaragdinum nigrum (p. 400), Lygosoma (Emoa) cyanogaster aruensis (p. 405) and Lygosoma (Hinulia) jobiense elegans (p. 397), were junior secondary homonyms at the time of description, respectively of Eumeces niger Jacquinot & Guichenot, 1853) (now Emoia nigra, see Brown, 1991), Eumeces aruensis Doria, 1874 (now Sphenomorphus jobiensis (Meyer, 1874), see Boulenger, 1887 and Zweifel, 1980), and Hinulia elegans Gray, 1838 (the current identity of which is unknown (Cogger et al., 1983), although from the illustration subsequently provided by Gray (1845), it may be a synonym of Eulamprus tenuis (Gray, 1831)). Lygosoma jobiense elegans is also a junior primary homonym of Lygosoma elegans Boulenger, 1897 (now Lobulia elegantoides (Ahl, 1925) and a junior secondary homonym of Euprepes (Tiliqua) elegans Fischer 1883 (now Lygosoma fernandi (Burton, 1836)) and itself a junior primary homonym of Euprepes elegans Peters, 1854, long considered to be part of the genus Mabuya Fitzinger, 1826, but more recently implicitly tranferred to the resurrected genus Euprepis Wagler (see Mausfeld et al., 2002), and still more recently to Trachylepis Fitzinger, 1843 (see Bauer, 2003) with recognition that Euprepis was not available for the Afro-Malagasy lineage formerly in Mabuya; Fischer (1884) proposed the replacement name Euprepes leoninus for Euprepes (Tiliqua) elegans.
Mertens (1929) provided the substitute name Dasia smaragdinum melas (now in Lamprolepis) for the secondary homonym L. smaragdinum nigrum Sternfeld, 1918, although the recognition of subspecies within Lamprolepis smaragdinum has mostly been avoided for over half a century (Kinghorn, 1928; Burt and Burt, 1932; Hediger, 1933, 1934; Greer, 1970; McCoy, 1980; Mys, 1988; Crombie & Pregill, 1999), and neither Sternfeld’s nigrum nor Mertens’ substitute name has subsequently appeared in the literature. Should the Nissan Atoll Lamprolepis be accorded taxonomic recognition in the future, Sternfeld’s nigrum is to be used as this name is no longer homonymous.
The homonymy of Sternfeld’s elegans was noted by Loveridge (1948: 346), although as no author since Sternfeld has applied the name to a species or subspecies distinct from Sphenomorphus jobiensis, no substitute name is needed.
An eighth skink taxon described by Sternfeld (1918) is also a homonym, though this time as a senior primary homonym, and this homonymy has not been previously recognised, nor is any subsequently published name available for the junior homonym. Sternfeld (1918) described Lygosoma (Emoa) cyanogaster keiensis, while Kopstein (1926) described Lygosoma (Homolepida) keiensis, both from the Kei Islands (now Kai Islands, or Kepulauan Kai, Indonesia). The epithet is formed incorrectly in both instances, as the name Lygosoma is neutral in gender. The epithet keiensis is to be corrected to keiense according to Articles 31.2 and 34.2 of the International Code of Zoological Nomenclature (International Commission on Zoological Nomenclature, 1999). This mandatory action has no further implications for the primary homonymy observed. Sternfeld’s subspecies is currently placed in the genus Emoia, while Kopstein’s species is currently placed in Sphenomorphus.
The purpose of this paper is to provide a replacement name for Kopstein’s junior homonym, and a more thorough description and diagnosis of the species, still known only from the holotype.
Lygosoma (Emoa) cyanogaster keiense Sternfeld, 1918
Lygosoma [(Emoa)] cyanogaster, Kopstein, 1926: 90.
Emoia cyanogaster, Loveridge, 1948: 366.
[Emoia] keiensis, Mittleman, 1952: 25.
Holotype.— SMF 15334 (Mertens, 1967: 74).
Type locality.— “Kei-Inseln, ? Molukken […] Langgur (Kei)” (Sternfeld, 1918: 405).
Current status.— Emoia longicauda (Macleay, 1877).
Discussion of citation history.— Kopstein (1926) was the first to comment on Sternfeld’s subspecies, stating that all of the Moluccan specimens he had collected agreed with Sternfeld’s diagnosis of L. c. aruense. No material from the Kai or Aru Islands was available to him, but nevertheless he remarked that the purported differences in coloration and pattern were untenable in view of the great chromatic variation within Lygosoma species. Consequently, he referred his specimens to a monotypic L. cyanogaster. Loveridge (1948) rejected both aruense and keiense as well, but erroneously stated that Sternfeld had not provided any diagnosis. Mittleman (1952) listed the species-group taxon keiense in combination with Emoia, but this action did not reflect an intent to elevate it to species status for he noted “[I]n no sense does this list purport to be a checklist of the forms considered valid.” Brown (1954) left the taxonomic status of the subspecies indeterminate, though restricting E. c. cyanogaster (Lesson, 1826) to the Solomon Islands, and recognising E. c. longicauda (Macleay, 1877) for populations in New Guinea. Mertens (1967) gave as current status Emoia cyanogaster keiensis without reference to a source, although presumably following Brown’s listing. Brown (1991) in his monographic revision of the genus, placed Sternfeld’s subspecies (misspelt twice as E. c. keinensis) in the synonymy of E. longicauda (Macleay, 1877), raising the latter to species status, although noting that further material was needed to resolve the status of the Aru and Kei Island populations. How et al. (1998), reporting on material from the Aru Islands, followed Brown’s (1991) synonymy of Sternfeld’s subspecies by not using trinominals in E. longicauda.
Sphenomorphus capitolythos, nomen novum for Lygosoma keiense Kopstein, 1926
Lygosoma [(Homolepida)] keiensis Kopstein, 1926: 86.
L[ygosoma]. (Sph[enomorphus].) keiensis, Brongersma, 1942b: 156.
[Sphenomorphus] keiensis, Greer & Parker, 1967: 19.
Holotype.— RMNH 5088, an ethanol-preserved specimen of undetermined sex, collected March 1923, by Dr Felix Kopstein.
Type locality.— “Elat, Gross-Kei” (Kopstein 1926: 86). In contemporary geographical nomenclature, this locality is Elat (or Banda Elat) on Kai Besar, Kepulauan Kai, Maluku Tenggara Province, Indonesia (5°39’S 132°39’E).
Diagnosis.— A medium-sized species of Sphenomorphus (SVL 81 mm) with short limbs, widely separated when adpressed. The combination of grooved subdigital lamellae, a scaly lower eyelid lacking a central window, four supraoculars, third pair of chin shields medially separated by three scales but in lateral contact with the infralabials, no postsupraocular scale, and a temporal region with no fragmentation or division of the last two supralabial scales, single primary temporal scale or upper and lower secondary temporal scales, and with the upper secondary temporal overlapping the lower secondary temporal, will differentiate this species from all other members of the Sphenomorphus group of lygosomine skinks (Greer, 1979a) in Indonesia, the New Guinea region, and Australia.
Description of holotype.—Nomenclature of head scales follows Taylor (1935), with exceptions as noted.
Head scalation.— Nasals moderately separated, nostril slightly posteroventrally located in nasal; supranasals and postnasals absent; prefrontals large, broadly separated medially; frontoparietals in broad contact; interparietal with parietal eye at junction of middle and posterior third; parietals in contact behind interparietal; nuchals 2/3; a single scale between upper secondary temporal and anterior nuchal on each side; supraoculars four, first two in contact with frontal; supraciliaries nine; posteriormost supraciliary (= anterior pretemporal of Greer, 1983) extends medially behind fourth supraocular, to contact frontoparietal on right side, but not on left side; posteriormost supraciliary not in contact with postocular, which instead wedges dorsally between penultimate supraciliary and uppermost postsubocular (= posterior pretemporal of Greer, 1983); single anterior and single posterior loreal, each with height and length similar, but posterior loreal dorsally a little longer than tall; presuboculars four; suboculars one; postsuboculars five; lower eyelid scaly, lacking a transparent window; supralabials seven, fifth below centre of eye, but separated from it by subocular series, sixth and seventh undivided; postsupralabials two; primary temporal single; secondary temporals two, lower overlapped by upper (on left side, the primary temporal is abnormally small, and the lack of its ventral part results in the last supralabial extending anteriorly to contact the postocular series; the arrangement of the scales of this region on the right side is typical of other sphenomorphine skinks with undivided temporal scales); external auditory meatus round, maximum diameter a little smaller than height of eye, tympanum deeply recessed; anterior margin of external auditory meatus lacking lobules; infralabials seven, first two in contact with postmental; three pairs of transversely enlarged chin chields, all laterally contacting the infralabial series; first pair of chin shields in medial contact; second pair separated by a median scale; third pair separated by three median scales.
Body and limb scalation.— Body scales in 31 rows at midbody (32 slightly anterior to this level); paravertebral scales not broader than adjacent dorsal scales, 59 from anteriormost nuchal to last scale anterior to level of hindlimbs (63 to level of posterior margin of hindlimbs); all scales polished and glossy; at high magnification, dorsal and lateral scales with many fine low longitudinal striations, but ventral scales smooth; median pair of preanals enlarged, overlapping more lateral preanals; lamellae below fourth toe 16, with a postaxial groove distinguishing a larger anterior and smaller posterior portion; scales above fourth toe six basally, three at middle of toe, reducing to two distally, with only terminal scale single; a sharp demarcation between rounded granular scales on sole and imbricate dorsal scales.
Measurements.— Snout-vent length 80.5 mm; axilla-groin interval 44 mm; tail distally regenerated; forelimb length 15 mm; hindlimb length 22 mm; head length 14.9 mm; head width 11.3 mm; head depth 8.8 mm.
Osteology (based on a radiograph of the type, and on direct examination of the dentition). — Presacral vertebrae 26, first three lacking ribs; sacral vertebrae two; tail regenerated from 17th postsacral vertebrae; ribs 6-8 sternal, 9-11 mesosternal, last four ribs short; phalangeal formula of manus and pes 184.108.40.206.3 and 220.127.116.11.4 respectively; premaxillary teeth nine (sum of left and right sides); maxillary teeth 18/17; postorbital bone possibly absent (the radiograph does not show a clearly distinct element in this region, although the postfrontal, jugal and squamosal are distinguishable).
Coloration.— Colour of the recently preserved specimen (after at most three years in preservative) was described by Kopstein (1926) as dorsally brown with irregularly arranged dark flecks, half a scale in size. Sides of the neck with faded grey reticulations. Lateral surfaces lighter brown. Ventral side uniform yellowish. The coloration in 2006, 80 years later, was similar, but paler. The dark flecks are located on both dorsal and lateral surfaces of body and tail, and are mostly located in the centre of scales, but occasionally obliquely oriented to cross scale rows. The upper and lower labial scales are dark-edged.
Kopstein remarked that his species was most closely related to L. unilineatum De Rooij, 1915. The latter taxon was recently reassigned to Eugongylus Fitzinger, 1843 by Greer and Shea (2000). However, S. capitolythos is not assignable to Eugongylus, based on supradigital scalation (multiple rows of scales in S. capitolythos vs. a single row in Eugongylus), nuchal scale orientation (obliquely to parietals vs. flush with parietals), preanal scales (median pair enlarged and overlapping adjacent lateral preanal scales vs all preanals more or less equal in size, with median pair overlapped by adjacent lateral preanal scales), number of presacral vertebrae (26 vs 28-30) and supranasals (absent vs usually present). In the first three of these character states, and in possessing nine premaxillary teeth, the species is clearly a member of the Sphenomorphus group of Greer (1979a). However, within this major lineage, the generic assignment of this species is less clear. In most scalational features, and in coloration, the species is similar to Glaphyromorphus nigricaudis (Macleay, 1877), from which it differs in having deeply grooved subdigital lamellae (vs smoothly rounded, ungrooved) and a greater number of paravertebral scales (63 vs 52-58, see Greer, 1979b). In the former feature, it is more similar to Sphenomorphus species occurring in New Guinea (Smith, 1937; Shea & Greer, 1999). However, it differs from the most speciose group in this region (the S. maindroni group, Greer & Shea, 2004) in lacking a postsupraocular scale, although in similarity with the postsupraocular of the S. maindroni group, the terminal supraciliary (or anterior pretemporal) extends to the frontoparietal (but on one side only).
There has been no rigorous morphological or biochemical analysis of relationships within the Sphenomorphus group of lygosomine skinks that has included both Glaphyromorphus and members of the S. maindroni group. The genus Sphenomorphus itself is currently undiagnosed in terms of derived character states within the Sphenomorphus group, and is likely to consist of more than one lineage on the basis of morphological variation. Similarly Glaphyromorphus is currently undiagnosed in terms of derived character states within the Sphenomorphus group, and represents a morphotype rather than a lineage (Greer, 1989, 1990). DNA sequence evidence for the polyphyly of Glaphyromorphus has been presented by Reeder (2003). Reeder also reported a major monophyletic lineage consisting of all Australian genera within the Sphenomorphus group, although his sampling of non-Australian representatives of the group was limited to just three species (representing Prasinohaema, Scincella and the Sphenomorphus muelleri species group). Hence, it is possible that his Australian lineage may include some non-Australian taxa, including some species currently assigned to Sphenomorphus. Given the present uncertainty about the limits and relationships of Sphenomorphus and Glaphyromorphus, we tentatively retain S. capitolythos in Sphenomorphus.
The specific epithet is a noun in apposition and honours Felix Kopstein (born 4 June 1893 in Vienna, Austria-Hungary [Adler stated Austria, but at that time Austria was part of the Habsburg Austro-Hungarian dual monarchy]; died 14 April 1939 in The Hague, the Netherlands), a physician and author of the last review of the Moluccan reptilian fauna (Adler, 1989). Capitolythos is derived from caput (Latin for ‘head’) and lythos (Greek for ‘stone’), in reference to the two probable German compound words in the name Kopstein, German for head and stone respectively. This unconventional form of patronym was chosen because of the existence of Lygosoma emigrans kopsteini Brongersma (1942a). The species emigrans is presently assigned to the genus Glaphyromorphus (Greer, 1990) but the subspecies has not been re-evaluated in subsequent studies of the genus in the Lesser Sundas (Auffenberg, 1980; Greer, 1990; Aplin et al., 1993), and has only been mentioned in a listing of Brongersma’s named taxa by Hoogmoed (1995).
Dr Marinus Hoogmoed (RMNH) kindly loaned the holotype of the species to the senior author for examination. Thanks are also extended to Mr Bertus van Tuijl, Dr Isaäc J.H. Isebrücker, Dr Axel Groenveld and Ms Elsbeth Zwart (ZMA) for granting access to the library and providing working space to the junior author. Dr Mieke Konings (Amsterdam) most kindly assisted in assessing the correctness of the name capitolythos.
Edited: L.P. van Ofwegen
Adler, K.K., 1989. Herpetologists of the past. In: Contributions to the history of herpetology (ed. K. Adler).— Society for the Study of Amphibians and Reptiles, Ithaca, New York, 5-141.
Ahl, E., 1925. Herpetologische Notizen.— Zoologischer Anzeiger 65: 18-20.
Aplin, K.P., R.A. How & Boeadi, 1993. A new species of the Glaphyromorphus isolepis species group (Lacer-tilia; Scincidae) from Sumba Island, Indonesia.— Records of the Western Australian Museum 16: 235-242.
Auffenberg, W., 1980. The herpetofauna of Komodo with notes on adjacent areas.— Bulletin of the Florida State Museum, Biological Sciences 25: 39-156.
Bauer, A.M., 2003. On the identity of Lacerta punctata Linnaeus 1758, the type species of the genus Euprepis Wagler, 1830, and the generic assignment of Afro-Malagasy skinks.— African Journal of Herpetology 52: 1-7.
Boulenger, G.A., 1887. Catalogue of lizards in the British Museum (Natural History). Vol. III. Lacertidae, Gerrhosauridae, Scincidae, Anelytropidae, Dibamidae, Chamaeleontidae: i-xii, 1-1-575, pl. I-XL.— British Museum (Natural History), London.
Boulenger, G.A., 1897. Descriptions of new lizards and frogs from Mount Victoria, Owen Stanley Range, New Guinea, collected by Mr A.S. Anthony.— Annals and Magazine of Natural History (6)19: 6-13, pl. I-II.
Brongersma, L.D., 1942a. Notes on scincid lizards.— Zoologische Mededeelingen 24: 125-152, pl. VI.
Brongersma, L.D., 1942b. On the arrangement of the scales on the dorsal surface of the digits in Lygosoma and allied genera.— Zoologische Mededeelingen 24: 153-158.
Brown, W.C., 1954. Notes on several lizards of the genus Emoia with descriptions of new species from the Solomon Islands.— Fieldiana (Zoology) 34: 263-276.
Brown, W.C., 1991. Lizards of the genus Emoia (Scincidae) with observations on their evolution and biogeography.— Memoirs of the California Academy of Sciences 15: 1-94.
Burt, C.E. & M.D. Burt., 1932. Herpetological results of the Whitney South Sea Expedition. VI. Pacific Island amphibians and reptiles in the collection of the American Museum of Natural History.— Bulletin of the American Museum of Natural History 63: 461-597.
Burton, E., 1836. [untitled].— Proceedings of the Zoological Society of London 1836: 62.
Cogger, H.G., E.E. Cameron & H.M. Cogger, 1983. Zoological Catalogue of Australia Volume 1. Amphibia and Reptilia: i-vi, 1-313.— Australian Government Publishing Service, Canberra.
Crombie, R.I. & G.K. Pregill, 1999. A checklist of the herpetofauna of the Palau Islands (Republic of Belau), Oceania.— Herpetological Monographs 13: 29-80.
Doria, G., 1874. Enumerazione dei rettili raccolti dal Dott. O. Beccari in Amboina, alle Isole Aru ed alle Isole Kei durante gli anni 1872-73.— Annali del Museo Civico di Storia Naturale di Genova 6: 325-357, pls. XI-XII.
Duméril, [A.]M.C. & [A.]M.A. Duméril, 1851. Catalogue méthodique de la collection des reptiles: 1-224.— Gide et Baudry, Paris.
Fischer, J.G., 1883. Beschreibung neuer Reptilien.— Jahresbericht über das Naturhistorische Museum zu Hamburg (Aus dem Osterprogramm des Akademischen Gymnasium) 1882: 1-16.
Fischer, J.G., 1884. Herpetologische Bemerkungen.— Abhandlungen aus dem Gebiete der Naturwissenschaften herausgegeben vom Naturwissenschaftlichen Verein in Hamburg 8: 1-11, pl. vii.
Fitzinger, L.I., 1826. Neue Classification der Reptilien nach ihren Natürlichen verwandtschaften. Nebst einer verwandtschafts- Tafel und einem verzeichnisse der Reptilien- Sammlung des K. K. Zoologischen Museum’s zu Wien: [i-v],1-128.— Verlage von J. G. Heubner, Wien.
Fitzinger, L., 1843. Systema reptilium. Fasciculus primus. Amblyglossae: 1-106, i-vi.— Braumüller et Seidel Bibliopolas, Vindobonae.
Gray, J.E., 1831. A synopsis of the species of the Class Reptilia. In: The Animal Kingdom arranged in conformity with its organization, by the Baron Cuvier, member of the Institute of France, &c. &c. &c. with additional descriptions of all the species hitherto named, and of many not before noticed. Volume 9. The class Reptilia arranged by the Baron Cuvier, with specific descriptions (ed. E.Griffith and E. Pidgeon). pp. 483-600.— Whittaker, Treacher, and Co., London.
Gray, J.E., 1838. Catalogue of the slender-tongued saurians, with descriptions of many new genera and species.— Annals and Magazine of Natural History (2)2: 287-293.
Gray, J.E., 1845. Reptiles. In: The Zoology of the Voyage of H.M.S. Erebus & Terror, under the command of Captain Sir James Clark Ross, R.N., F.R.S., during the years 1839 to 1843. Volume 2 (ed. J. Richardson & J.E. Gray). pp. 1-8, pl. 1-4, 8-9, 12-14, 20.— London: Longman, Brown, Green and Longmans.
Greer, A.E., 1970. The relationships of the skinks referred to the genus Dasia.— Breviora 348: 1-30.
Greer, A.E., 1974. The generic relationships of the scincid lizard genus Leiolopisma and its relatives.— Australian Journal of Zoology, Supplementary Series 13: 1-67.
Greer, A.E., 1979a. A phylogenetic subdivision of Australian skinks.— Records of the Australian Museum 32: 339-371.
Greer, A.E., 1979b. A new Sphenomorphus (Lacertilia: Scincidae) from the rainforests of northeastern Queensland.— Records of the Australian Museum 32: 373-382.
Greer, A.E., 1983. A new species of Lerista from Groote Eylandt and the Sir Edward Pellew Group in Northern Australia.— Journal of Herpetology 17: 48-53.
Greer, A.E., 1989. The biology and evolution of Australian lizards: i-xvi, 1-264.— Chipping Norton, Australia: Surrey Beatty & Sons.
Greer, A.E., 1990. The Glaphyromorphus isolepis species group (Lacertilia: Scincidae): diagnosis of the taxon and description of a new species from Timor.— Journal of Herpetology 24: 372-377.
Greer, A.E. & F. Parker, 1967. A new scincid lizard from the northern Solomon Islands.— Breviora 275: 1-20.
Greer, A.E. & G. Shea, 2000. A phylogenetically important lygosomine skink resurrected from taxonomic obscurity: Lygosoma unilineatum de Rooij, 1915.— Journal of Herpetology 34: 85-91.
Greer, A.E. & G. Shea, 2004. A new character within the taxonomically difficult Sphenomorphus group of lygosomine skinks, with a description of a new species from New Guinea.— Journal of Herpetology 38: 79-87.
Günther, A., 1872. Notice of a new species of lizard (Eumeces albofasciolatus) from north Australia.— Annals and Magazine of Natural History (4)10: 370-371.
Hediger, H., 1933. Über die von Herrn Dr A. Bühler auf der Admiralitäts-Gruppe und einigen benachbarten Inseln gesammelten Reptilien und Amphibien.— Verhandlungen der Naturforschenden Gesellschaft in Basel 44: 28-52.
Hediger, H., 1934. Beitrag zur Herpetologie und Zoogeographie New Britanniens und einiger umliegender Gebiete.— Zoologische Jahrbücher. Abteilung für Systematik, Ökologie und Geographie der Tiere 65: 441-585.
Hoogmoed, M.S., 1995. In memoriam Prof. Dr Leo Daniel Brongersma (1907-1994).— Zoologische Mededelingen 69: 177-201.
How, R.A., B. Durrant, L.A. Smith & N. Saleh, 1998. Emoia (Reptilia: Scincidae) from the Banda Arc islands of eastern Indonesia: variation in morphology and description of a new species.— Records of the Western Australian Museum 19: 131-139.
International Commission on Zoological Nomenclature, 1999. International Code of Zoological Nomenclature, Fourth Edition.— The International Trust for Zoological Nomenclature, London.
Jacquinot, H. & A.Guichenot, 1853. Reptiles et poissons. In: Zoologie. Tome 3 (ed. M. Hombron and H. Jacquinot). pp. 1-56. In: Voyage au Pôle Sud et dans l’Océanie sur les corvettes “l’Astrolabe” et “la Zélée” exécuté par ordre du Roi pendant les années 1837-1840 (ed. J. Dumont d’Urville).— Paris.
Kinghorn, J.R., 1928. Herpetology of the Solomon Islands.— Records of the Australian Museum 16: 123-178, pl. XIII-XV.
Kopstein, F., 1926. Reptilien von den Molukken und den benachbarten Inseln.— Zoologische Mededee-lingen 9: 71-112.
Lesson, R-P., 1826. Reptile. In: Atlas de Zoologie.Voyage autour de monde, exécuté par ordre du Roi, sur la corvette de sa Majesté La Coquille, pendant les années 1822-1825, Pl. 3-4.— Arthus Bertrand, Paris.
Loveridge, A., 1948. New Guinean reptiles and amphibians in the Museum of Comparative Zoölogy and United States National Museum.— Bulletin of the Museum of Comparative Zoology 101: 305-430.
Macleay, W., 1877. The lizards of the “Chevert” expedition.— Proceedings of the Linnean Society of New South Wales 2: 60-69.
Mausfeld, P. & A. Schmitz, 2003. Molecular phylogeny, intraspecific variation and speciation of the Asian scincid genus Eutropis Fitzinger, 1843 (Squamata: Reptilien: Scincidae): taxonomic and biogeographic implications.— Organisms Diversity & Evolution 3: 161-171.
Mausfeld, P., A. Schmitz, W. Böhme, B. Misof, D. Vrcibradk & C.F.D. Rocha, 2002. Phylogenetic affinities of Mabuya atlantica Schmidt, 1945, endemic to the Atlantic Ocean archipelago of Fernando de Noronha (Brazil): necessity of partitioning the genus Mabuya Fitzinger, 1826 (Scincidae: Lygosominae).— Zoologische Anzeiger 241: 281-293.
McCoy, M., 1980. Reptiles of the Solomon Islands. Handbook (7): i-vi, 1-80.— Wau Ecology Institute, Wau, Papua New Guinea.
Mertens, R., 1922. Verzeichnis der Typen in der herpetologischen Sammlung des Senckenbergischen Museums.— Senckenbergiana 4: 162-183.
Mertens, R., 1924. Herpetologische Mitteilungen.— Senckenbergiana 6: 177-185.
Mertens, R., 1929. Die Rassen des Smaragdskinkes, Dasia smaragdinum Lesson.— Zoologischer Anzeiger 84: 209-220.
Mertens, R. 1967. Die herpetologische Sektion des Natur-Museums und Forschungs-Institutes Senckenberg in Frankfurt a.M. nebst einem Verzeichnis ihrer Typen.— Senckenbergiana biologica 48 (Sonderheft A): 1-106.
Meyer, A.B., 1874. Übersicht der von mir auf Neu-Guinea und den Inseln Jobi, Mysore und Mafoor im Jahre 1873 gesammelten Amphibien.— Monatsberichte der königlichen Akademie der Wissenschaften zu Berlin 1874: 128-140.
Mittleman, M.B., 1952. A generic synopsis of the lizards of the subfamily Lygosominae.— Smithsonian Miscellaneous Collections 117: 1-35.
Monk, K.A., Y. De Fretes & G. Reksodiharjo-Lilley, 1997. The Ecology of Nusa Tenggara and Maluku. The Ecology of Indonesia Series, Vol. V: i-xvii, 1-966.— Periplus Editions, Hong Kong.
Mys, B., 1988. The zoogeography of the scincid lizards from North Papua New Guinea (Reptilia: Scincidae). I. The distribution of the species.— Bulletin de l’Institut Royal des Sciences Naturelles de Belgique 58: 127-183.
Peters, W.C.H., 1854. Diagnosen neuer Batrachier, welche zusammen mit der früher (24. Juli und 17. August) gegebenen Übersicht der Schlangen und Eidechsen mitgetheilt werden.— Bericht über die zur Bekantmachung geeigneten Verhandlungen der königlich-preussischen Akademie der Wissenschaften zu Berlin 1854: 614-628.
Reeder, T., 2003. A phylogeny of the Australian Sphenomorphus group (Scincidae: Squamata) and the phylogenetic placement of the crocodile skinks (Tribolonotus): Bayesian approaches to assessing congruence and obtaining confidence in maximum likelihood inferred relationships.— Molecular Phylogenetics and Evolution 27: 384-397.
Rooij, N. de, 1915. The Reptiles of the Indo-Australian Archipelago. I. Lacertilia, Chelonia, Emydosauria: i-xiv, 1-384.— Leiden: E.J. Brill.
Scott, F., F. Parker & J.I. Menzies, 1977. A checklist of the amphibians and reptiles of Papua New Guinea.— Wildlife in Papua New Guinea 77/3: 1-18.
Shea, G.M. & A.E. Greer, 1999. Two senior synonyms and a name change for the New Guinea skink Sphenomorphus stickeli (Loveridge, 1948).— Journal of Herpetology 33: 136-141.
Smith, M.A., 1937. A review of the genus Lygosoma (Scincidae: Reptilia) and its allies.— Records of the Indian Museum 39: 213-234.
Sternfeld, R., 1918. Zur Tiergeographie Papuasiens und der pazifischen Inselwelt.— Abhandlungen der Senckenbergischen naturforschenden Gesellschaft 36: 375-436.
Taylor, E.H., 1935. A taxonomic study of the cosmopolitan scincoid lizards of the genus Eumeces with an account of the distribution and relationships of its species.— University of Kansas Science Bulletin 23: 1-643.
Tyler, M.J., 1968. Papuan hylid frogs of the genus Hyla.— Zoologische Verhandelingen 96: 1-203, pl. 1-4.
Welch, K.R.G., P.S. Cooke & A.S. Wright, 1990. Lizards of the Orient: A Checklist: i-v, 1-162.— Malabar: Robert E. Kreiger Publishing Co.
Whitaker, R., Z. Whitaker & D. Mills, 1982. Reptiles of Papua New Guinea.— Wildlife in Papua New Guinea 82/2: 1-53.
Zweifel, R.G., 1979. Variation in the scincid lizard Lipinia noctua and notes on other Lipinia from the New Guinea region.— American Museum Novitates 2676: 1-21.
Zweifel, R.G., 1980. Results of the Archbold Expeditions. No. 103. Frogs and lizards from the Huon Peninsula, Papua New Guinea.— Bulletin of the American Museum of Natural History 165: 387-434.