Stellio' in Herpetology and a Comment on the Nomenclature and Taxonomy of Agamids of the Genus Agama (Sensu Lato) (Squamata: Sauria: Agamidae)

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HERPETOZOA 8 (1/2): 3 - 9 Wien, 30. Juli 1995

A brief review of the origin and use of 'stellio' in herpetology and a comment on the nomenclature and taxonomy of agamids of the genus Agama (sensu lato) (Squamata: Sauria: Agamidae)

Kurzübersicht über die Herkunft und Verwendung von "stellio" in der Herpetologie und Kommentar zur Nomenklatur und Taxonomie von Agamen, Gattung Agama (sensu lato) (Squamata: Sauria: Agamidae)

KLAUS HENLE

ABSTRACT The name 'stellio' has received a wide and variable application in herpetology. Its use antedates modern nomenclature. The name was applied mainly to various agamid and gekkonid species. In modern usage, confusion exists primarily with its use as a genus, i. e., Siellio. To solve this problem, STEJNEGER (1936) desig- nated Siellio saxatilis of LAURENTI, 1768 which is based on a figure in SEBA (1734) as the type species. This species is unidentifiable. In an unpublished thesis, MOODY (1980) split the genus Agama (s. 1.) into six genera. He overlooked STEJNEGER's (1936) designation and reused Stellio for the stellio-group of agamid lizards. Many authors followed MOODY (1980). Recently, some authors pointed out that Siellio is unavailable but did not fully discuss the implications for agamid nomenclature. It is argued that a satisfactory nomenclature is difficult with current knowledge of agamid taxonomy. It is suggested to restrict Laudakia to L. tuberculata and to use Ploce- denna for the stellio-group (sensu stricto).

KURZFASSUNG Der Name "stellio" sah eine breite und vielfältige Verwendung in der Herpetologie, die weit über die moderne Nomenklatur zurückreicht. Vorwiegend wurden mit diesem Namen verschiedene Gecko-Arten und Agamen der Gattung Agama (sensu lato) bezeichnet. Verwirrung entstand bezüglich der korrekten Verwendung als Gattungsname. Zur Lösung der Probleme designierte STEJNEGER (1936) Stellio saxatilis LAURENTI, 1768 als Typusart, ein auf einer Abbildung von SEBA (1734) basierendes, nicht identifizierbares Taxon. In einer unveröffentlichten Doktorarbeit unterteilte MOODY (1980) die Gattung Agama (s. 1.) in sechs eigenständige Gattungen. Er übersah STEJNEGER's (1936) Festlegung der Species typica und brachte Stellio wieder in Gebrauch für Agamen der stellio-Gruppe. Viele Autoren folgten ihm. Nur vereinzelt haben Autoren darauf hingewiesen, daß Siellio nicht verfügbar ist, aber die Konsequenzen für die Nomenklatur von Agamen unvollständig diskutiert. Es wird aufgezeigt, daß beim gegenwärtigen Wissensstand der Taxonomie von Aga- men eine befriedigende Nomenklatur schweirig ist. Als Vorschlag wird unterbreitet, den Gebrauch des Gat- tungsnamens Laudakia auf L. tuberculata zu beschränken und den Namen Plocedenna für die stellio-Gruppe im engeren Sinne zu verwenden.

KEYWORDS Stellio: availabilty, type species; Agama, Laudakia, Plocedenna: nomenclature, taxonomy, phylogeny

THE USE OF 'STELLIO' IN HERPETOLOGY

The name 'stellio' is one of the oldest ANANJEVA & al. 1990; CORBETT 1990; vernacular names known for any lizard. It JOGER 1991). Few authors differ: AR- has received a widespread and varying ap- NOLD (1986) retains the genus Agama plication both in pre-Linnean and post-Lin- (sensu lato) but recognized Stellio as a sub- nean times. Currently, most authors use genus. BAIG & BÖHME (1991) also use the generic name Stellio for agamids of the Agama (s. 1.); they point out that Stellio is stellio-group of the formerly more inclu- not available as a generic name. LEVITON sive genus Agama (e. g., SCHÄTTI 1989; & al. (1992) also briefly state that Stellio ©Österreichische Gesellschaft für Herpetologie e.V., Wien, Austria, download unter www.biologiezentrum.at

K. HENLE

is unavailable but suggest to call agamids According to ARISTODELES, the of the stellio-group Laudakia. They are animals in question which he called followed by others (e. g., FRANZEN & 'ascalabotes' live primarily in Thrace and SCHMIDTLER 1993). As none of these Nikandro but also in Sicily. He, and PLI- authors explain in detail why the name is NIUS, regard their biology as considerably unavailable and only partly discuss the en- different in the two areas; e. g., they be- suing consequences, it is of general lieve that Greek animals are harmless but interest to briefly review the use of the Italian ones are venomous. The mentioned name stellio. The more so, as BAIG & Greek area is outside the current distribu- BÖHME (1991) and LEVITON & al. tion of T. mauritanica (RIEPPEL 1981) (1992) arrive at different conclusions. and all other geckos except of Cyrtopodion The name 'stellio' has a Latin origin kotschyi (see BÖHME 1981). It is partly and appeared for the first time in the Latin covered by the range of Agama stellio literature around the first century a. d. (BEUTLER 1981). In Sicily, only geckos (BYL 1986, 1987). Most etymologists are found. Above arguments demonstrate agree that the animals in question were that geckos must have been included in the named 'stellio' because of their colour name 'stellio1, and that it is likely that pattern resembling stars, though other ex- agamids were included as well. planations exist (e. g., SCHNEIDER The continued confusion of geckos [1811] believes that 'stellio' alludes to and agamids into the post-Linnean time dorsal tubercles in addition to white spots; adds support for this hypothesis: e. g., RA- see also BYL [1987]). Philologists discuss FINESQUE-SCHMALTZ (1810) names T. at length the possible idendity of the ani- mauritanica from Segesta, Sicilly, Agama mals and arrive at different conclusions. scarpina. Nevertheless, it is very likely Whereas some argue that scorpions are the that PLINIUS is talking mainly about appropriate candidates others regard lizards geckos as he knows that ' stelliones' eat and generally geckos as the correct animals their own slough and informs us that the (see LEITNER 1972; BYL 1987 - and lit- slough was used as a remedy against erature cited therein). Etymological expla- epilepsy. He also writes that the animals nations are generally neglected in attempts are found mainly on doors and windows of to identify the animals in question. buildings and in graveyards - common Most philologists use PLINIUS as habitats of T. mauritanica in southern Eu- the prime source of information about rope (HENLE - pers. observ.); only grave- 'stellio' as do later zoologists (e. g., GES- yards would be possible habitats of A. NER 1554; PAOLI 1771; CETTI 1777; stellio. Also, PLINIUS lived predomi- SCHNEIDER 1811; BONAPARTE 1835; nantly in southern Italy and thus his per- DUMÉRIL & BIBRON 1836). These zo- sonal experience must relate mainly to ologists agree that PLINIUS called geckos T. mauritanica. 'stellio' while the Greek folk used the Confusion surrounding the correct name 'ascalabotes' for these reptiles. Most use of stellio in modern nomenclature of these zoologists, though not all, also starts right from the beginning. LIN- believe that the species in question is Ta- NAEUS (1758) uses stellio as the specific rentola mauritanica. Their arguments are epithet of his Lacerta stellio. While there based on the use of the name 'stellio' in never has been any doubt that his L. stellio concurrent vernacular language. Latins and refers to the European agamid species A. people in many places in Toscany called [s. 1.] stellio, trouble begins with LAU- geckos 'stellio'. Rural Sards called geckos RENTI's (1768) introduction of Stellio as •pistilloni* (CETTI 1777). Currently, 'lo a generic name. He includes eight species stellione' (and 'lo scarpino') is still used in his genus Stellio all based on figures by some Sicilians for T. mauritanica and descriptions of SEBA (1734). Creating (HENLE - unpubl. pers. experience). Stellio, LAURENTI deviated from his However, elsewhere in Italy, geckos usu- usual habit in the creation of new genera ally were called tarentola. by not using a specific name introduced by ©Österreichische Gesellschaft für Herpetologie e.V., Wien, Austria, download unter www.biologiezentrum.at

Stellio, agamid taxonomy and nomenclature

LINNAEUS as a new generic name. In- to accommodate agamids of the stellio- stead, he omits L. stellio from his treatise. group. MOODY (1980) neither reviews This certainly fooled many modern au- the nomenclature of this or any of the thors. other five genera into which he splits LAURENTI (1768) does not desig- Agama s. I. nor formally diagnosed his nate a type species, and because of the lack genus Stellio. He and authors following of tautonymy, none could be fixed auto- him overlook STEJNEGER's type desig- matically. Of the eight species included in nation. Because of the continuing use of Stellio, one is a skink, 3 or 4 are varanids Stellio and the difficult accessability of (opinions differ on this point), and the re- SEBA (1734), it seems to be advisable to maining 3 to 4 remain unidentifiable (see include SEBA's description together with a STEJNEGER 1936). reprint of his figure (fig. 1): Stellio is commonly used by zoolo- 'Tecoixin, seu Lacerta saxatilis, spi- gists of the 18th and 19th century but nosa, cauda crassula. never in the sense of LAURENTI. Some Squamulae dilute cinereae, ex russo argue that the name Stellio applies to obumbratae, spinis horrent albicantibus, geckos and use it for various genera (e. g., quarum quaelibet è macula nigro-fusca, Gekko, Uroplatus) but do not include any tanquam basi, porrigitur. Neque spinis ca- of the species of LAURENTI's genus ret ipsum caput, nee femora. Cauda cras- Stellio. SCHNEIDER (1792, 1811) is the sula squamulis tenuibus, cinereo-luteis, prime advocate of this opinion (see above tantum vestitur. Quini sunt pedum postico- for further authors). In contrast to rum digiti, anticorum quaterni. In locis sa- SCHNEIDER, LATREILLE (in SONNINI xosis Americae degit.' & LATREILLE 1801), DAUDIN (1802), As can be seen from this description MERREM (1820), and WAGLER (1830) and the accompanying figure (fig. 1), the apply Stellio to various agamids (mainly name cannot be attributed to any known Agama s. 1. but also Uromastyx). They species with certainty. Returning to my also do not include any of LAURENTI's earlier argument, I regard it as very likely (1768) taxa. GRAY (1825) even derives a that SEBA (1734) also combined geckos family name from Stellio and includes and agamids in his Lacerta saxatilis. various agamid, iguanid, and cordylid gen- Noteworthy, the figure is much more styl- era in his Stellionidae. MERREM (1820) ized than most others in SEBA (1734), makes matters worse by substituting Gecko making it likely that he and/or the artist stellio for Tare/itola mauritanica and did not have a reference individual avail- Agama tetradactyla for LAURENTI's able. The colouration could well be that of Stellio saxatilis. Fortunately, his action did A. stellio though it may also apply to some not receive any wide application. For un- species of Gekko. The general body form known reasons, the use of Stellio for and the shape of the regenerated tail also geckos is abandoned in the second half of could make this genus a candidate for La- the 19th century, but remains in use for certa saxatilis, particularly, if one consid- various agamids. It is applied primarily to ers that in the Dutch version of SEBA, agamids belonging to the genus Agama s. four claws instead of four digits are men- 1., particularly to species in the stellio- tioned (KLAVER, pers. comm.). How- group. ever, the toes definitely exclude geckos of Because of the confusion surrounding the genus Gekko. In any case, an unequiv- the name Stellio and to stabilize well-es- ocal species identification is impossible. tablished names (Varanus or Eumeces), Importantly, the depressed body and the STEJNEGER (in SMITH 1933) designates regenerated tail definitely excludes aga- the taxon Stellio saxatilis as the type mids of the genus Agama s. 1. Thus, Stel- species which he believes to be unidentifi- lio must be regarded as a nomen dubium able. With STEJNEGER's action, Stellio and remains certainly unavailable for the falls into disuse until MOODY (1980) res- A. stellio-group. urrects the genus in an unpublished thesis E o 1 2 3 8 9 10 11 12 13 14 15 16 17 18

Fig. 1: Tecoixin, seu Lacerta saxatilis of SEBA (1734: plate 79, fig. 4): type species of the genus Stellio. Reproduced with permission of the Landesbibliothek Stuttgart. Abb. 1: SEBAs (1734: Tafel 79, Figur 4) Tecoixin, seu Lacerta saxatilis: Species typica der Gattung Stellio. Wiedergabe mit Genehmigung der Landesbibliothek Stuttgart. ©Österreichische Gesellschaft für Herpetologie e.V., Wien, Austria, download unter www.biologiezentrum.at ©Österreichische Gesellschaft für Herpetologie e.V., Wien, Austria, download unter www.biologiezentrum.at

Siellio, againid taxonomy and nomenclature

IMPLICATIONS FOR AG AMID NOMENCLATURE

Two possible consequences for the taxonomic units as genera, as has been nomenclature of Western Palearctic agam- done by most later authors. ids follow from above conclusion: either Finding a new generic name for the retaining all species groups within the stellio-group, and some of the other genus Agama s. 1., or, if one prefers to el- groups, causes considerable problems. The evate them to full genera, one has to find next oldest names, Cyclosaurus WAGLER another name for the stellio-group. None in MICHAHELLES 1833 (based on A. of the two approaches is completely satis- hispida) and Saura WAGLER in MICHA- factory in view of our current knowledge HELLES 1833 (based on A. agama + A. of the phylogeny of agamids. ANANJEV A atra), are synonyms of Agama s. str. The & SOKOLOVA (1990) and JOGER (1991) same holds true for Podorrhoa FITZIN- provide biochemical evidence that Phryno- GER 1843 (based on A. agama) and Psam- cephalus is more closely related to the morhoa FITZINGER 1843 (based on A. stellio-group than the latter is to the muta- aculeata). Eremioplanis FITZINGER bilis-group (but see MOODY's 1980 phy- 1843 (based on A. mutabilis), Trapeloides logram). Therefore, Phrynocephalus should FITZINGER 1843 (based on A. sanguino- be incorporated in Agama s. 1., if one ac- lenta), and Planodes FITZINGER 1843 cepts the first approach. Furthermore, the (based on A. agilis) all are synonyms of first approach would conceal the advances Trapelus. Acanthocercus FITZINGER gained in understanding the phylogeny of 1843 is based on Stellio cyanogaster, a agamids. member of the African clade. This clade, Discussing the problems of the sec- however, certainly is not congeneric with ond approach, we first have to summarize the Palaearctic clade (JOGER 1991). Ac- briefly the results of MOODY (1980). cording to JOGER's phylogenetic tree, it MOODY splits the formerly more inclu- may be regarded as congeneric with Pseu- sive genus Agama into six species groups dotrapelus FITZINGER 1843 (both de- which he elevates to genus level: Agama, scribed on the same page) or as a sister Trapelus, Pseudotrapelus, Stellio, Xen- genus to the latter. agama, and Brachysaura (his group VI). The next available name within Within this group, the first three taxa are Agama s. 1., Laudakia GRAY 1845, has characterized by a derived karyotype (2n been used by LEVITON & al. (1992) for = 44-48 chromosomes) while Stellio re- the stellio-group. The type species of Lau- tains the ancestral karyotype (2n = 36 dakia, A. tuberculata, is a member of the chromosomes). Xenagama and Brachysau- Palearctic clade and may be used for the ra have not yet been examined. Trapelus is stellio-group (sensu lato). However, prob- further characterized by a small and deeply lems remain with this approach as L. tu- sunken tympanum forming an external au- berculata differs considerably in hemi- ditory tube-like meatus, and Pseudotra- penial characters (apomorphic characters: pelus by a swollen tail base due to the divided hemipenes, forked sulcus, and presence of keratinized coarse scales. Xen- transversal division of the apex) from the agama is unique in having the traverse •s?e///o-group (sensu stricto) which is closer processes of the first caudal vertebrae pro- to Agama (sensu restricto) (BÖHME jecting perpendicular to the longitudinal 1988). Furthermore, it has a derived kary- axis (versus projecting slightly posteriorly otype (2n = 34 versus 2n = 36 - ancestral in all others). Stellio is the only genus karyotype of the stellio-group sensu having the derived characters of several stricto) (GORMAN 1973) lacking one pair spinous scales located immediately behind of microchromosomes. Although the kary- the orbit and/or in the region between the otype of L. tuberculata should be reexam- ear as well as a nuchal crest. Thus, using inecl as microchromosomes may be over- the data presented by MOODY (1980), it looked (GORMAN 1973), it should be appears sufficiently justified to accept his pointed out that Pseudotrapelus is also ©Österreichische Gesellschaft für Herpetologie e.V., Wien, Austria, download unter www.biologiezentrum.at

K. HENLE

Table 1: New generic allocation of species formerly included in 'Siellio'. Tabelle 1: Neue Gattlingszuordnung für Arten, die seither zu 'Stellio' gerechnet wurden.

Plocedenna Plocedenna/Laudakia Laudakia Acanthocercus incertae sedis

melanura agroensis luberculala cyanogaster caucasia badakshana adramitanus eryihrogastra chemovi anneclens himalayana lännanensis atricollis lehmanni mi croie pi s phillipsii nupta nuiistanica yemenensis siellio pakisianica zonura sacra sioliczkana tarimensis

characterized by 22 micrichromosomes is a synapomorphy or is due to convergence (GORMAN & SHOCHAT 1972). Finally, (BÖHME 1988). based on albumin antisera and isozymes, Until the discussed phylogenetic the genus Phrynocephalus is a sister group problems are resolved by studying addi- of the stellio-group (sensu stricto) (JOGER tional species and by combining morpho- 1991). Unfortunately, JOGER (1991) had logical, karyological, and biochemical ap- no data available on L. tuberculata, and its proaches, I regard it as the best solution to relative phylogenetic position to stellio restrict the use of Laudakia to L. tubercu- (sensu stricto) and to Phrynocephalus re- lata. This approach leaves open the ques- mains unknown. Phrynocephalus also has tion of a genus name for the stellio-gvowp. a derived karyotype (2n = 46 - 48) (BÖH- Plocedenna BLYTH 1854 (based on A. ME 1981) hk& Agama, Trapelus, and Pseu- melanura) is the oldest available name for dotrapelus but different from Laudakia. In the stellio-group (sensu stricto) and should Phrynocephalus helioscopus persicus, the be used for it. Species for which it is not only species within the genus for which the known whether they are closer related to hemipenes are known, they also are highly the stellio-group or to L. tuberculata may derived (divided, forked sulcus, large ca- be referred to either cf. Laudakia or cf. lyces). However, it remains unresolved Plocedenna (see tab. 1 for a list of spec- whether the similarities between Phrynoce- ies). phalus and L. tuberculata (and Trapelus)

ACKNOWLEDGMENTS I thank Dr. W. BÖHME (Bonn) for helpful uable suggestions and an introduction to the relevant discussions of the nomenclature and taxonomy of philological literature. geckos and agamids, and Dr. L. BODSON for val-

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Stellio, agamid taxonomy and nomenclature

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DATE OF SUBMISSION: September 13th, 1994 Corresponding editor: Werner Mayer

AUTHOR: Dr. Klaus HENLE, Projektbereich Naturnahe Landschaften, Umweltforschungszentrum Leipzig-Halle G. m. b. H, Permoserstraße 15, D-04318 Leipzig, Germany.