when the printing plates, which had been lost for two decades, were found. Pallas's description is accurate, 
mentioning the number of scutes on the shell, the number of toes on the manus and pes, and the color and shape of 
the shell; however, without his figure, T. ibera could not be recognized as a subspecies of T. graeca. 
Perhaps the most colorful contributor to tortoise nomenclature was Constantine Samuel Rafinesque. In 
1832, he began publishing The Atlantic Journal and Friend of Knowledge, which ceased publication two years later, 
after, among other things, making an enemy of Richard Harlan (Rafinesque, 1836:333). In this publication, 
Rafinesque (1832) provided an extremely brief description of Gopherus, specifically referring to the flattened nails of 
the manus. 
The second volume of Duméril and Bibron's encyclopedic Erpetologie Generale (1835) contained descriptions 
of Testudo [= Geochelone] nigrita and a new genus Homopus. T. nigrita was characterized as black with a crescent- 
shaped anterior carapacial edge, a single supracaudal, slightly imbricate scales on the forearm, and no cervical. 
Homopus was characterized by four digits on all limbs, ungulate feet and a solid carapace without a hinge. T. 
areolatus and T. signatus were referred to Homopus; however, H. signatus usually has five toes on the manus. 
In 1835, Fitzinger coined many of the generic and subgeneric names employed today. Geochelone, for 
example, was proposed by Fitzinger. Cylindraspis, the name for the extinct tortoises from Mauritius, Rodrigues and 
the Mascerenes, was delineated from other Geochelone by its single gular scute, which is only present in some 
Cylindraspis, and by another ambiguous character, lateral shell margin arcuate. Chelonoidis and Psammobates were 
also differentiated by ambiguous characters. Both Cylindraspis and Chelonoidis were proposed as subgenera of 
Geochelone, whereas Psammobates was considered a subgenus of Testudo. 
From 1838 to 1849 (see Waterhouse, 1880, for publication dates), Andrew Smith published pieces of his 
Illustrations of the Zoology of South Africa.... In the1839 installment, he illustrated and described two tortoises: 
Testudo verroxti [= Psammobates tentorius verroxii] and Testudo semiserrata [= Psammobates oculifer]; both defined 
primarily by coloration and shell shape. In 1840, Schlegel and Miiller described two new species, Testudo [= 
Manouria] emys and Testudo [= Indotestudo] forstenii. A good illustration was provided for T. emys, but T. 
forstenii was described only in the text of a footnote. Nevertheless, the written descriptions of both species are 
accurate, matching well the surviving types (Hoogmoed and Crumly, 1984). Four years later, Gray (1844) named 
Testudo horsfieldii from an "India, Affghanistan" [sic] juvenile. Gray's description did not include features sufficient 
to diagnose this species. He noted blunt hip spines, long gular scutes, large anal scutes and an oblong shell. 
Although well done for its time, Gray's descriptions make Schlegel and Miiller's contribution all the more 
impressive. 
1850-1900 
Duméril, Bibron and Duméril (1851) proposed the substitute name Testudo [= Manouria] emydoides for 
Testudo emys Schlegel and Miller. The Dumérils' description emphasized cervical shape, the shape and number of 
marginal scutes, the scales covering the limbs, and hip and tail spines. Twelve plastral scutes were noted; although 
they did not mention whether the pectoral scutes met medially or were separated by humero-abdominal contact. 
Because they did not mention the unusual condition in Manouria emys [ssp. emys], it seems likely that the pectoral 
scutes did meet on the midline. Because 25 marginals were noted, it is probable that the supracaudal was split. 
Another synonym of Geochelone emys was proposed a year later by Gray (1852), who provided a new genus name 
Manouria for his species, M. fusca. Manouria was characterized by a divided supracaudal, pectoral scutes not 
meeting on the midline and centrally located carapacial areolae. Gray believed that Manouria was an emydid but 
noted many similarities shared with testudinids. He characterized Manouria fusca using shell color, recurved anterior 
and posterior shell margin, and projecting gular region. 
In 1853, Edward Blyth recognized two new species: Testudo phayeri [= Manouria emys phayeri] and Testudo 
[= Indotestudo] elongata. Testudo phayeri was distinguished by its flat carapace, double supracaudal, unusual limb 
scalation and color. This name has been considered a synonym of G. emys, although the color is somewhat different 
from previously described G. emys. Roger Bour (pers. comm.) and I consider T. phayeri the senior synonym of G. 
nutapundi Nutaphand (1979). It seems likely that Manouria emys comprises two subspecies intergrading along a 
north-south cline. T. elongata was recognized by its elongated carapace and characteristic coloration. But, Blyth did 
not notice that the humero-pectoral sulcus transversely crosses the entoplastron, a feature that I consider diagnostic 
for Indotestudo. 
