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Neotype of Testudo terrestris Forsskål, 1775 (Testudines, Testudinidae)
JARMO PERÄLÄ1 AND ROGER BOUR2
1

Department of Biological and Environmental Sciences, PO Box 65 (Biocenter 3, Viikinkaari 1),

FIN-00014 University of Helsinki, Finland; E-mail: jarmo.perala@helsinki.fi
2

Laboratoire des Reptiles et Amphibiens, Muséum national d’Histoire naturelle,

25 rue Cuvier, 75005 Paris, France; E-mail: bour@mnhn.fr
Abstract. - We discuss and clarify the nomenclatural status of Testudo terrestris Forsskål, 1775. Testudo terrestris, ataxon based on a syntype series, was until now a valid name for tortoises from parts of Egypt, Arabia and the Levant,according to the International Code of Zoological Nomenclature. On the contrary, all type locality restrictions fromthe 20th century regarding the name Testudo terrestris Forsskål, done without the fixation of a name-bearing specimen(lectotype or neotype), are invalid. Because the name is valid, and because the syntypes, listed but not identified in
the original description, are untraceable, and to permanently fix the name and its type locality, we designate a neotype
for Testudo terrestris Forsskål based on a specimen from Aleppo, Syria.
Key words. - Taxonomy, Testudines, Testudinidae, Testudo terrestris Forsskål, 1775, nomenclatural validity, neotype.
Forsskål and Mediterranean tortoises. - During his
fatal travel (1761-1763) in the eastern Mediterraneancountries, the Finn Petter Forsskål (1732-1763) encountered tortoises, briefly described by the name Testudo
terrestris in his diary, which was published as a posthumous work by Carsten Niebuhr (1733-1815), a German
traveller and surveyor, in 1775 [= Forsskål, 1775].
Strauch (1862: 69) remarks in a footnote that Forsskål
(1775) discusses a tortoise, Testudo terrestris, which
occurs in Aleppo and Lebanon, without describing it,
and Anderson (1896: 68) writes that “it is uselessattempting to identify this animal” “from Loheia, northof Hodeida,” with reference to Forsskål (1775).
Forsskål’s itinerary took him through the following
localities: Malta, Smyrna (Izmir), Constantinople, Alexandria, Rosetta, Cairo, Suez, Jedda, ending up in Yarim,
Arabia Felix (Yemen), where Forsskål died of malaria in
1763. Tortoises were reported from Al Lu ayyah(Yemen), Cairo (Egypt), Lebanon, Lattakia (Al L dhiq
- yah) and Aleppo (both in Syria) (Fig. 1). Forsskål
(1775) indicated that the chelonians are called “Zolhafae” by the Arabs (Forsskål, 1775); as for their distribution, he specified: “Kahirae non frequens vivit,
Aleppo autem & ad Libanon copiosor,” translated by
Daudin (1801: 225) as: “It is rare in Cairo, but can befound rather abundantly near Aleppo and towards
Mount Lebanon”. At best, Forsskål himself was thus
able to see tortoises in situ in northern Egypt (Nile Delta
and Suez area) only, that is those which are actuallycalled either Testudo kleinmanni Lortet, 1883 (the Egyptian tortoise) or the vicariant T. werneri Perälä, 2001.
This interpretation is in accordance with the vernacular© 2004 by Asiatic Herpetological Research

name of the Egyptian tortoise, fide Anderson (1898: 30;“Sohlafa” pronounced “Zihlifa”) and Flower (1933:
742; “Educated people in Egypt employ the word ‘Zalheefah’”). Niebuhr’s subsequent route back home took
him via Basra, Baghdad, Mosul, Diyarbak1r, Aleppo,
Lebanon and Palestine (Hansen, 1962; Niebuhr, 1772,1973). Niebuhr, the only surviving member on the expedition to Arabia Felix, continued to Bombay (Mumbai),
India, from where he shipped the natural history notes(Niebuhr, 1772: xix) and specimens (Niebuhr, 1973:
120) collected by Forsskål via London to Copenhagen.
A considerable amount of material in the Forsskål collection in Copenhagen was destroyed in bombings by
the British in 1807 (Nielsen, 1993). The type of T. terrestris is said to be "apparently lost" (Webb in Iverson,1992). The name is not fixed to a single name-bearing
specimen.
Nomenclatural validity of Testudo terrestris. - Testudo
terrestris Forsskål, 1775, one of the oldest species groupnames in the genus Testudo Linnaeus, 1758 sensu lato(in the sense of Lapparent de Broin, 2001; Perälä
2002a), is nomenclaturally valid, according to thepresent International Code of Zoological Nomenclature(ICZN, 1999): it was published before 1931 and thus
does not need to have a full diagnosis by which it can be
identified. Only “a description or a definition” per se isneeded (ICZN, 1999: Article 12). In this respect theremark by Forsskål (1775) pertaining to the tortoises’
length (“...foot-long...”), as well as that made of the
shapes of plastra of both sexes, is enough, even if thesecharacters were incorrect. Moreover, although the nameVol.10, p. 115

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Figure 1. Map of geographical localities mentioned in
Forsskål's (1775) description, which together constituted
the type locality of Testudo terrestris Forsskål until thepresent work.

is accompanied by the remark “Obs.” (Forsskål, 1775:
12), it is also listed under the headings “Descripta” (p.
viii) as well as “Nominata” (p. ix) alongside with valid
names such as Testudo triunguis Forsskål, 1775, that is,current Trionyx triunguis. The book contains a multitudeof other descriptions of valid taxa which cannot berejected based on the arguments (see below) about theuse of the Latin language, a contemporary standard,alone. Based on Niebuhr’s subsequent route back home,
Gasperetti et al. (1993) suggested that it may actually be
Niebuhr who is responsible for the name T. terrestris
Forsskål. As Niebuhr’s role is not explicit in the original
publication, the authorship of the name remains with
Forsskål (ICZN, 1999: Art. 50.1.1).
Testudo terrestris overlooked, rejected, then revalidated. - After its publication, and during about 180
years, the nominal species Testudo terrestris Forsskål,1775, although nomenclaturally valid, was neverthelessoverlooked by most authors (except Strauch, 1862 andAnderson, 1896), some of them (Gray, 1831; Duméril

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and Bibron, 1835) only mentioning Testudo zolkafa andTestudo zohalfa, respectively, based on Forsskål’s(1775) vernacular Zolhafae, and which nomina nuda
appear in the above works as synonyms of Testudo
graeca Linnaeus, 1758 and Testudo mauritanica
Duméril and Bibron, 1835, respectively. On the other
hand, at least one scientist worked to revive the nominal
species. A brief history of the case is presented by Bourin David (1994: 86). The name Testudo terrestris
Forsskål was resurrected, and therefore revalidated, by
Wermuth (1956: 402), who improperly (without specimen fixation) designated “Arabia” as type locality.
Simultaneously, Wermuth considered to ask the International Commission on Zoological Nomenclature toinvalidate the nominal species Testudo terrestris
Forsskål; unfortunately, his attitude changed radically(1958: 149-153) and he used this name to designate the
Near Eastern population of Testudo (as Testudo graeca
terrestris Forsskål), restricting the type locality to“Libanon-Gebirge, Israel” [sic], and mistakenly extending its range to Libya. It must be outlined that there is a
gap in the range of Testudo graeca (sensu lato) complex
tortoises between Israel and Libya, the latter containing
the range of the recently described species Testudo
cyrenaica Pieh & Perälä, 2002. Wermuth had at his disposal one Libyan specimen (SMF 36127; paratype of T.
cyrenaica), from Dernah, which he thought to be identical with Middle-Eastern tortoises (Pieh & Perälä, 2002).
Wermuth’s validation was disapproved by Buskirk (inErnst et al., 2000; unpublished manuscript from theearly 1990s; and pers. comm. to both authors); Highfield (in Ernst et al., 2000; and: http: //www.tortoisetrust.org/articles/newfloweri.html); J. F. Parham(pers. comm.); and Perälä (1996); among others. Their
opposition is based on several arguments such as: adescription is lacking with reference to the remark“Obs.” in Forsskål (1775); the species cannot be identified from the “description”; there is no type specimen; a“false” type locality; the name “Testudo terrestris” isjust Latin for a terrestrial chelonian, used without intention to describe a new species.
Need of a neotype. - Rather to resurrect Testudo terrestris Forsskål (Wermuth, 1956), it would have been preferable: (1) either to suppress this imprecise name (andwhich is a more recent homonym of the nominal species
Testudo terrestris Fermin, 1765, for which Wermuth had
to successfully request the invalidation by the International Commission on Zoological Nomenclature; ICZN,
1963: Opinion 660); (2) either to use it to name Testudo
kleinmanni, a name revalidated only in the 1950s by
Mertens & Wermuth (1955), and independently by Loveridge & Williams (1957). It could still be possible toask the ICZN to officially suppress the name Testudo

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Figure 2. Neotype of Testudo terrestris Forsskål, 1775, specimen n° NMW 18674: 2, sub-adult female. Upper left: lateral view (right side); Upper right: dorsal view; Lower left: ventral view. Lower right: living specimen of Testudo terrestris, adult female, from Aleppo (topotype); CL = 185 mm, MI = 143 mm, HE = 99 mm.

terrestris Forsskål, 1775. However, we reject this option
because: (1) the name is presently widely used (in all
recent check-lists, with more than fifty references),
although with vagueness about the identity of the concerned population; (2) there is no valid name to designate the species of Testudo living in the Middle-East,
more precisely in the area of the upper Euphrates Tigris drainage; (3) the name Testudo terrestris
Forsskål, 1775 became available by the very ruling of
the International Commission on Zoological Nomenclature (ICZN, 1963), and it is unlikely that the Commission would reverse its opinion. Accepting thenomenclature proposed by Wermuth, one of us (RB)proposed to emend the type locality of Testudo terrestris
Forsskål to the vicinity of Aleppo (= Halab), Syria(Bour, 1989: 14), in the interest of clarifying the statusof this taxon and as a first step towards the descriptionof a neotype. However, such restriction of the type
locality, as well as the earlier restrictions proposed by
Wermuth (1956, 1958), are invalid according to theCode (ICZN, 1999; Art. 76.3), because these actions

were not done in connection with the selection of a lectotype or neotype. Therefore we here propose the formal
description of a neotype of Testudo terrestris Forsskål,1775. All the animals listed (although not identified;therefore untraceable) by Forsskål (from Al-Lu ayyah,
Cairo, Lebanon, Lattakia and Aleppo) actually represent
syntypes, according to the Code (ICZN, 1999; Art.
72.1.1), and thus the type locality of T. terrestris encompasses the region containing all those localities (ICZN,
1999; Art. 73.2.3). A neotype could legitimately be
selected from any of those per se. Besides sea turtles,
only Centrochelys sulcata (Miller, 1779), a land tortoise(possibly introduced), and Pelomedusa subrufa(Lacepède, 1788), a fresh-water turtle, are known tooccur in Yemen (Obst & Wranik, 1987; Gasperetti et al.,
1993; Al Safadi, 1997); therefore no Testudo sp. could
have been observed in Al-Lu ayyah by the Forsskål Niebuhr expedition. Following the earlier choice, madein accordance with the available data, and also with thecurrent taxonomical practice (e.g., Perälä, 2002b), in
order to preserve the stability of the nomenclature, andVol.10, p. 117

Asiatic Herpetological Researchto objectively delimit Testudo terrestris Forsskål fromall other species in the Testudo graeca (s.l.) complex,
we choose a specimen collected in Aleppo (and which
locality has by chance a historical background in tortoise literature: cf. Siebenrock, 1913).
The neotype of Testudo terrestris. - To fix the name andtype locality, we hereby designate a specimen from theVienna Natural History Museum No. NMW 18674:2,
collected in Aleppo by Viktor Pietschmann in March
1910, as the neotype of Testudo terrestris Forsskål,1775, according to Articles 75 and 76 (ICZN, 1999).
The neotype is a subadult female with a straight-line
carapace length (CL) of 136.8 millimeters. Note: allmorphometric characters are according to the standards
published in Perälä (2001). As a result of our neotype
designation (ICZN, 1999; Art. 76.3), the type locality ofTestudo terrestris Forsskål, 1775 is restricted to Aleppo(Alep, Halab; 36°12’ N, 37°09’ E), Syria (Syrian Arab
Republic). (For the range of T. terrestris, see Perälä,
2002b.)
Description. – Sub-adult female. Most scutes marked
with about a dozen of conspicuous growth ridges andgrooves; areolae of the carapace feebly bumped, neatly
displaced caudally, and also dorsally on the costals.
Longitudinal profile high, regularly domed, highest at
third vertebral, slightly behind the middle of the shell(Fig. 2, top left). The outline of the shell short, squarish,
anterior and posterior free borders only very feebly cut
out. Vertebrals wide in dorsal view, the fourth the smallest; the first one with almost straight lateral borders,moderately wider anteriorly than posteriorly (Fig. 2, top
right). Cervical (= nuchal) four-sided, very wide; supracaudal distally as wide as vertebrals, regularly convex inits middle; all common sutures of marginals nearly sub
equal (= height of lateral ones, and length of anterior
and posterior ones); only marginals 9-11 are slightly
flaring. Gulars well prominent but short and narrow,their common suture very short; a pair of rather large
axillaries on each side; pectorals with a very short common (medial) suture, included about four times in the
medial suture of humerals; inguinals small, separated ina larger distal part and a very small proximal part contacting femorals; rear lobe of plastron hardly mobile,
short, anals both long and wide, with parallel anterior
and posterior borders: their particular shape at first sightresembles the usual shape observed in males (Fig. 2,
bottom left). Head covered above with two large androughly pentagonal scutes, the frontal and the prefrontal, symmetrical about their common suture. Five nails
at each hand, the inner one smaller but well developed;
four at each foot. Anterior side of the fore-arm covered
between the elbow and the wrist by about fifteen large

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scutes, the four largest being triangular, neatly distinct
from the background of the scaly skin; outer border ofthis area covered by a row of six triangular, overlapping
scutes. Tail relatively long and regularly tapering, also
giving a rather masculine appearance, ending withslightly enlarged but discrete flat scutes; a small isolated
spur on each thigh.
General color greenish yellow, often lighter close to
the sutures, with large darker, grayish areas apparently
deep in the scutes. Blackish marks reduced on the carapace, limited to incomplete and irregular narrow lines
along the anterior sutures of the scutes (marginals, costals), also on the lateral borders of vertebrals; areolae or
areolar areas slightly and irregularly flecked with black,on costals 1-3 and on vertebrals 1-3. Dark patches wider
on the plastron, issued from the areolae, roughly extending along the rear third of each scute (from pectorals toanals), restricted to a narrow band on the humerals; limits of the patches are inconspicuous, with a gradualshading, delimiting few lighter or darker radiating lines.
Soft parts mostly yellowish, in places with a brownish
tinge, with well contrasting blackish flecks on the horny
beak (upper and lower, forming a ‘moustache’), on prefrontal, on the large triangular scutes of the fore-arms,
and on the heels; all nails are also heavily pigmented,
from dark brown to black.
Additional morphometric data derived from theneotype are presented in the following (all measurements in mm): maximum plastron length (PL) = 122.7,
midline plastron length (PL-m) = 111.8, maximum midbody width (MI) = 100.2, maximum width of shell at
posterior marginals (MA) = 104.4, maximum gularscute length (GU-l) = 15.3, maximum gular scute width(GU-w) = 27.3, gular scute height (GU-h) = 13.2, maximum shell height (HE) = 70.0, maximum width of anterior shell opening (ASO-w) = 69.9, maximum height ofanterior shell opening (ASO-h) = 21.9, left minimum
bridge length (BR) = 59.9, maximum humeral scutewidth (HUM-w) = 63.7, maximum pectoral scute width(PEC-w) = 86.6, maximum abdominal scute width(ABD-w) = 89.9, maximum femoral scute width (FEMw) = 64.1, maximum anal scute width (AN-w) = 51.9,
maximum nuchal scute length (NU-l) = 11.4, maximum
nuchal scute width (NU-w) = 10.0, intergular length(GU-m) = 13.4, interhumeral length (HUM-m) = 22.8,
interpectoral length (PEC-m) = 7.7, interabdominal
length (ABD-m) = 42.1, interfemoral length (FEM-m) =10.8, interanal length (AN-m) = 18.7, maximum widthof first vertebral scute (V1-w) = 35.2, maximum widthof second vertebral scute (V2-w) = 36.4, maximum
width of third vertebral scute (V3-w) = 41.0, maximum
width of fourth vertebral scute (V4-w) = 33.8, maximum width of fifth vertebral scute (V5-w) = 41.3, maximum length of first vertebral scute (V1-l) = 26.9,

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Asiatic Herpetological Researchmaximum length of second vertebral scute (V2-l) =28.0, maximum length of third vertebral scute (V3-l) =25.6, maximum length of fourth vertebral scute (V4-l) =23.3, maximum length of fifth vertebral scute (V5-l) =31.8, first costal length (C1) = 43.0, second costal length(C2) = 28.7, third costal length (C3) = 28.1, fourth costal length (C4) = 22.9, maximum dorsal width of supracaudal (SUP-d) = 23.4, maximum ventral width ofsupracaudal (SUP-v) = 40.4, maximum median lengthof supracaudal (SUP-l) = 21.7, maximum head width(HEAD) = 21.6, minimum distance between right eye
and tympanum (EYE-TY) = 6.9, minimum distance
between right eye and nostril (EYE-NO) = 6.5.
Figure 2 (lower right) depicts a living female (topotype) Testudo terrestris Forsskål from the type locality,
Aleppo, Syria.

Acknowledgments

Vol.10, p. 118

David, P. 1994. Liste des reptiles actuels du monde. I.
Chelonii. Dumerilia 1:1-127.
Duméril, A. M. C. and G. Bibron. 1835. Erpétologie
générale ou histoire naturelle complète des reptiles.
Tome 2. Librairie Encyclopédique de Roret, Paris.
i-ii, 680 pp.
Ernst, C. H., R. G. M. Altenburg and R. W. Barbour
2000. Turtles of the World: World Biodiversity
Database CD-ROM Series. Expert Center for Taxonomic Identification, University of Amsterdam &UNESCO-Publishing, Paris.
Flower, S. S. 1933. Notes on the recent reptiles andamphibians of Egypt, with a list of the species
recorded from that Kingdom. Proceedings of the
Zoological Society, London 1933:734-851.

We warmly thank James Buskirk (Oakland, CA) andJames Parham (University of California, Berkeley) for
personal communications, as well as two anonymous
reviewers for comments, which had a positive impact on
the manuscript, Richard Gemel, Heinz Grillitsch andFranz Tiedemann (all from the Naturhistorisches
Museum, Vienna) for the loan of specimens, the Academic Kippis Society (AKS) for spiritual guidance, andAnnemarie Ohler (Muséum national d’Histoire
naturelle, Paris) for logistical help.

Forskål [sic], P. 1775. Descriptiones Animalium,
Avium, Amphibiorum, Piscium, Insectorum, Vermium; quae in itinere orientali observavit Petrus
Forskål Prof. Haun. Post mortem auctoris edidit
Carsten Niebuhr. Heineck & Faber, Hauniae(Copenhagen). xxxiv, 164 pp.

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