A NOMENCLATURAL HISTORY OF TORTOISES(Family Testudinidae) Charles R. CrumlyDepartment of HerpetologySan Diego Naturtal History Museum SMITHSONIANHERPETOLOGICAL INFORMATIONSERVICENO. 751988 SMITHSONIANHERPETOLOGICALINFORMATIONSERVICE The SHIS series publishes and distributestranslations, bibliographies, indices, and similaritems judged useful to individuals interested inthe biology of amphibians and reptiles, butunlikely to be published in the normal technicaljournals. Single copies are distributed free tointerested individuals. Libraries, herpetologicalassociations, and research laboratories are invitedto exchange their publications with us.We wish to encourage individuals to share theirbibliographies, translations, etc. with otherherpetologists through the SHIS series. If youhave such items please contact George Zug forinstructions. Contributors receive 50 free copies.Please address all reguests for copies andinguiries to George Zug, Division of Amphibians andReptiles, National Museum of Natural History,Smithsonian Institution, Washington, D.C. 20560,U.S.A. Please include a se 1 f -addressed mailinglabel with reguests. The cover illustration is a reproduction of the plate from the originaldescription of John Edward Gray (1870a). IntroductionPrerequisite to most revisionary studies is an examination of the nomenclatural history of the taxa understudy. This is often a gruelling task, but is rarely published. Thus, each subsequent revisor must repeat the task ofgathering all the references together and extracting the pertinent information. The following idiosyncratic reviewsummarizes my view on the nomenclatural history of tortoises (family Testudinidae).Most, but not all, of the synonyms noted by Wermuth and Mertens (1977) are discussed below. The genericnames in brackets are the currently used names (Loveridge and Williams, 1957; Auffenberg, 1974) or my recentlyrecommended usage (Crumly, 1983, 1984b, 1984c). I have chosen not to detail the nomenclatural history of theIndian Ocean tortoises or the Galapagos tortoises (= Geochelone elephantopus complex of Crumly, 1984c) becausetlieir histories are too complex and because many biological questions need to be considered in order to interpret theirhistories. Detailed historical discussions of die tortoises of the Indian Ocean islands and the Galapagos can be foundelsewhere (Van Denburgh, 1914; Slevin, 1959; Bour, 1984). Furthermore, the nomenclatural history of entirelyextinct groups of tortoises (e.g. Siylemys, Cylindraspis) are not discussed.1750-1800Linnaeus (1758) proposed eleven turtle binomials in his 10th edition of Sysiema Naturae; two of these arenow in the family Testudinidae. Both tortoises had been recognized earlier. He referred to Edwards's (1751, p. 204)figure in the description of Testudo graeca, and his description of Testudo [= Psanunobaies] geometricus referred toseveral earlier publications. T. graeca was characterized by elephantine feet with 4 toes on the pes. Psaminobatesgeometricus was distinguished by the slightly conical carapacial scutes and the radiating shell pattern. Wallin(1977) demonstrated that Linnaeus based his description of/', geometricus on a specimen of Geochelone elegans aswell as other descriptions. To preserve common usage, Hoogmoed and Crumly (1984) selected a lectotype one ofthe earlier descriptions (= syntypes) employed by Linnaeus. In the 12th edition o^ Systema Naturae, Linnaeus(1766) recognized Testudo [= Geochelone] dcnticulata by its eroded carapacial marginal (now known to be moreprominent in Geocfielone carbonaria, fide Williams, 1960) and the elephantine feet.Carsten Niebuhr edited Petrus Forskal's notes on his Middle Eastern travels (Forskal, 1775). These notesincluded Arabic names for all the animals he saw and the description of Testudo terresiris [= T. graeca terrestris] plussome life history and distributional notes. Forskal felt that Arabic names would assist others in future fieldwork;however, if "Libanon" had not been mentioned there would have been no means of recognizing T. g. terrestris; it isthe only tortoise living here. Miller's description (1780) of Testudo [= Geochelone] sulcata is based solely on anillustration. The illustration itself is unlabeled. However, Miller's index of illustrations, which includes the name T.sulcata, enabled Loveridge and Williams (1957) to verify his authorship. Carl Peter Thunberg, a Linnaeuscorrespondent and collector coined Testudo [= Homopus] areolata (Thunberg, 1787). He noted the flattened mid-dorsal region of the carapace, the large imbricate scales of the forearm, and the depressed carapacial areolae. Twoyears later, Gmehn (1789), in Linnaeus's 13th edition o{ Systema Naturae, named Testudo hermanni. Gmelin notedtwo especially distinctive features: 1) The "caudae apice unguiculato" [an apical tail spine] and 2) "marginisscutella 24" [24 marginal scutes indicating a divided supracaudal].Schoepffs beautifully illustrated volume (1792b) included descriptions of two new species, Testudomarginata and Testudo [= Geochelone] elegans. The marginated tortoise was delineated by its expanded and flaredposterior carapacial margin. The shell pattern and less conical carapacial scutes distinguished Testudo [= Geochelone]elegans from P. geometricus. With the close of the 18th century, seven of the approximately 40 living species oftestudinid were recognized. 1800-1850Shaw's description (1802) of Testudo [= Geochelone] radiata was the first new tortoise named in the 19thcentury. He relied heavily on Grew's earlier description (1681, Musaeum Regalis Socieiatis). Shaw recognized thatthis species was not synonymous with T. [= Psammobaies] geometrica. By virtue of its larger size and flattened,rather than conical, carapacial scutes, it was distinct from the earlier known T. geometrica. In France, Daudinpublished the second volume of llistoire Naturelle des Reptiles (1802) and described Testudo [= Gopherus]polyphemus and Testudo luteola [= Psammobates geometricus]. Testudo [= Gopherus] polyphemus wasdistinguished by its small scales, flattened forelimbs and ash-gray color. The description of T. luteola notes themany similarities shared with P. geometrica but distinguished it by less elevated conical carapacial scutes. Subsequently (Loveridge & Williams 1957; Muller, 1939), the heights of carapacial scutes have been shown to beextremely variable .The parade of new names for tortoises continued with August Schweigger's (1812) famous Prodromusmonographiae Cheloniorum. The Prodromus included three new tortoises, Testudo [= Kinixys] erosa, Testudo [=Chersina] angulata and Testudo [= Geochelone] gigantea. Testudo erosa was distinguished by shell shape and itsdenticulate margin. The number of marginal scutes and the elongation of the anterior periphery of the shell was alsomentioned, but not the unique carapacial hinge. Schweigger used the type locality from Shaw's earlier description(1802) of T. denticulata (preoccupied by T. denticulata Linnaeus, 1766); the locality, "America septentrionali," wasin error. T. angulata was distinguished by shell shape, shell color, and narrow anterior marginals. Because elevenplastral scutes were mentioned the presence of a single gular scute can be inferred. The provenance of this specieswas unknown to Schweigger, who had examined two living specimens at the Paris museum. The description of T.gigantea is quite difficult to interpret; over the years it has come to refer to the tortoise now living on the island ofAldabra. However, three very different positions have been articulated (Bour, 1982b, 1984b; Crumly, 1986;Pritchard, 1986).In 1820, Kuhl named Testudo [= Psammobates] ocuUfera and characterized it by its color pattern and did notcompare it to any other species. In the same year, Merrem authored his Versuch eines Systems der Amphibien(1820). He introduced a new generic name, Chersine, for all tortoises. This name subsumed many turtles besidestestudinids (for example, Kinosternon scorpioides, Clemmys muhlenbergi). In part, this name has been considered asynonym of Testudo.Four years later, Spix (1824) named Testudo [= Geochelone] carbonaria, and three species now consideredsynonyms of G. denticulata, i. e., Testudo hercules, T. sculpta and T. cagado. Hoogmoed and Gruber (1983)discovered that, except for 4 specimens of T. sculpta in the Zoologische Sammlung des Bayerischen Staates (= ZSM)collections (ZSM 2753/0 a, b, c and 2738/0), Spix's tortoise types had been destroyed near the end of Worid War II.Spix used color, elevated conical carapacial scutes and eroded lateral shell margins in his description of T. carbonarla.In 1827, a banner year for new tortoise taxa, four species and 2 genera were described. Richard Harian (1827)named Testudo [= Geochelone] elephantopus based on its reflected anterior shell margin and absent terminal tailspine. Thomas Bell (1827) named three new species and two new genera. Bell distinguished Pyxis arachnoides, anew species and genus, from other tortoises on the basis of its unique anterior plastral hinge. He coined Kinixys forthose tortoises with posteriorly mobile carapacial lobe and recognized two species, Kinixys castenea and K.homeana. K. castenea was later synonymized with K. erosa by Gray (1831b). Bell did not mention and wasapparendy unaware of Schweigger's (1812) earlier description of 7. [= Kinixys] erosa.In the following year. Bell (1828) named three more taxa: Testudo [= Psammobates] tentoria, Testudo [=Geochelone] pardalis and Testudo actinoides. Testudo actinoides was synonymized with Testudo [= Geochelone]elegans by Boulenger (1889). T. tentorius was distinguished from other tortoises on the basis of conical carapacescutes, a radiating carapacial color pattern and unequal sized anterior marginals. This last trait was not mentioned byearlier workers, and apparently Bell considered it a very significant diagnostic character. Bell diagnosed T. [=Geochelone] pardalis by the light-golden yellow colored carapace with black flecks, the superior location of thepleural areolae, and the equal sized anterior marginals, emphasizing the pleural areolae trait.John Edward Gray's Cataphracta volume of the Synopsis Reptilium appeared in 1831; he named two newtaxa. Kinixys belliana was described in the Additions and Corrections chapter and was distinguished from otherKinixys species by the shape of the carapace, cervical scute and shell margin. He believed that this new species wasintermediate between K. erosa and A^. denticulata (?= Testudo denticulata Shaw 1802). During this same year. Gray(1831a) coined Chersina (not the same as Merrem's Chersine of 1820) for Testudo angulata Schweigger. He notedeleven plastral scutes (from which I infer that there was a single gular scute) as the distinguishing feature of thisSouth African species.Peter Simon Pallas (1831) named Testudo ibera [= T. graeca ibera], in the third volume of a posthumouslypublished three volume set. The original date of publication for this third volume has been variously cited: 1814 byWermuth (1958), Darevsky and Mertens (1973), and Wermuth and Mertens (1961, 1977), and 1827 by Mertens(1946). Neither is correct. In Pallas's Zoographica Rosso-Asiatica, the description of Testudo ibera is speciesnumber 14 on page 18, thus 1814. Zaunick (1925) noted that the third volume of the set was published in 1831 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 ofthe 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. In1832, 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 ofthe manus.The second volume of Dumeril and Bibron's encyclopedic Erpetologie Generale (1835) contained descriptionsof 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, forexample, was proposed by Fitzinger. Cylindraspis, the name for the extinct tortoises from Mauritius, Rodrigues andthe Mascerenes, was delineated from other Geochelone by its single gular scute, which is only present in someCylindraspis, and by another ambiguous character, lateral shell margin arcuate. Chelonoidis and Psammobates werealso differentiated by ambiguous characters. Both Cylindraspis and Chelonoidis were proposed as subgenera ofGeochelone, whereas Psammobates was considered a subgenus of Testudo.From 1838 to 1849 (see Waterhouse, 1880, for publication dates), Andrew Smith published pieces of hisIllustrations of the Zoology of South Africa.... In thel839 installment, he illustrated and described two tortoises:Testudo verroxii [= Psammobates tentorius verroxii] and Testudo semiserrata [= Psammobates oculifer]; both definedprimarily 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 areaccurate, matching well the surviving types (Hoogmoed and Crumly, 1984). Four years later. Gray (1844) namedTestudo horsfieldii from an "India, Affghanistan" [sic] juvenile. Gray's description did not include features sufficientto 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 Muller's contribution all the moreimpressive. 1850-1900Dumeril, Bibron and Dumeril (1851) proposed the substitute name Testudo [= Manouria] emydoides forTestudo emys Schlegel and Miiller. The Dumerils' description emphasized cervical shape, the shape and number ofmarginal scutes, the scales covering the limbs, and hip and tail spines. Twelve plastral scutes were noted; althoughthey 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 pectoralscutes 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 nameManouria for his species, M. fusca. Manouria was characterized by a divided supracaudal, pectoral scutes notmeeting on the midline and centrally located carapacial areolae. Gray believed that Manouria was an emydid butnoted many similarities shared with testudinids. He characterized Manouria fusca using shell color, recurved anteriorand 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 limbscalation and color. This name has been considered a synonym of G. emys, although the color is somewhat differentfrom 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 anorth-south dine. T. elongata was recognized by its elongated carapace and characteristic coloration. But, Blyth didnot notice that the humero-pectoral sulcus transversely crosses the entoplastron, a feature that I consider diagnosticfor Indotestudo. John Leconte (1854) described Teleopus luxatus, a synonym of Geochelone emys. LeConte noted a splitsupracaudal scute and described in great detail the vertebral and marginal scutes. He also noted the tricuspid upperjaw and bicuspid lower jaw and further remarked that this species shared features with both testudinids and emydids.Louis Agassiz published Contributions to the Natural History of the United Stales of America in 1857. Agreat proportion of this work (Parts 11 & 111 of three parts) dealt with turtles. Therein, Agassiz named and allocatedtwo species to a new genus Xerobates [= Gopherus]. It was diagnosed by a median premaxillary ridge, broadflattened unguligrade forelimbs, and a broad head covered with mostly small scales. Xerobates carolinus [=polyphemus] was not characterized, but distribution and life history notes were included. X. berlandieri was describedand distinguished by light yellow color in the middle of and rinmiing some carapacial scutes, by divergent gulars,and by small size as suggested by retained granular areolae of the shell. The syntypes of Xerobates berlandieri (twospecimens bearing the same number, USNM 60) are a juvenile and a hatchling, which explains the presence ofgranular areolae. These areolae are worn smooth in adults.Six years passed before any more currently valid names for tortoise species were proposed. Blyth (1863) thennamed Testudo [= Geochelone] platynota based on its large size, flat back, less numerous radiations on the carapacialscutes, and broader and lower carapacial shape. John Edward Gray (1863) "provisionally" named Kinixys spekii froman "imperfect" specimen from central Africa. The oblong shell, areolar color and lack of a carapacial hinge were usedby Gray to characterize this form. Loveridge and Williams (1957) tentatively synonymized K. spekii with K.belliana; this decision was later reversed by Broadley (1982). John G. Cooper (1863) described Xerobates [=Gopherus] agassizii in honor of Alexander Agassiz. Cooper inadequately distinguished X. agassizii from otherXerobates by shell shape and color. Four years later, Grandidier (1867) named a tortoise from Madagascar, Testudo[= Acinixys] planicauda. He characterized this species on the basis of color, a single supracaudal, a small nuchal, anda uniquely flattened tail.One of the few studies to deal specifically with the cranial osteology of land tortoises appeared in 1869. Inthis work, John Gray described two taxa, Testudo (Scapia)falconeri [= Manouria emys] and Peltastes (an invalidgenus including five species now allocated to at least three genera). Gray's characterization of T.falconeri detailedcranial structures that are not diagnostic, but the accompaning figure clearly illustrated features unique to M. emys;broad skull arches, a deep lower jaw, a narrow but long skull and the absence of an interdigitating surangular process.Gray further noted the similarities between Manouria fusca, another synonym of M. emys, and T. falconeri. Theinterruption of the alveolar surface by enlarged internal narial openings and slight prominences at the labial maxillo-premaxillary suture characterized Peltastes Gray (1869). Of the five species allocated to Peltastes by Gray, only P.[= Indotestudo] elongata was well characterized.A year later. Gray (1870a) named Testudo [= Geochelone] chilensis with only a very sketchy description,which wrongly contended that a nuchal plate [= cervical scute] is present. A plate of the presumed type is presentedby Gray, and the cervical scute is absent on the illustrated specimen. In that same year. Gray (1870b) publishedanother even shorter note that reviewed previous papers concerning the synonymy of T. chilensis. Also in 1870,Sclater noted that T. chilensis does not occur in Chile and that this species had been known to previous students.Fueling the fires of mutual dislike, he suggested a substitute name, Testudo argentina, which was never adopted.Gray published his Appendix to his earlier catalogue of British Museum turtles in 1872 and created another synonymof Geochelone, Centrochelys (type species Geochelone sulcata), defined by an absent cervical and large hip spines.Following Gray's flurry of activity, Giinther (1875) briefly reported on giant land tortoises in the BritishMuseum and named thiee new taxa from the Galapagos, Testudo ephippium, Testudo microphyes and Testudovicina. Gunther's descriptions are very good and include data on shells, skulls and limbs. Unfortunately, the lack ofgood locality data for referred material diminishes the utility of Gunther's descriptions. In fact, he incorrectlyidentified the island of origin for both Geochelone ephippium and Geochelone microphyes, whereas he was unable todiscover an exact island of origin to Geochelone vicina. Two years later, Gunther's monograph (1877) on BritishMuseum giant tortoises appeared. Therein, he named Testudo [= Geochelone] abingdonii. This monograph issecond only to Van Denburgh's (1914) classic study and exceeded even Van Denburgh in the quality illustration andin his discussion of non-shell material. Giinther (1877) thought the skulls and other post-cranial non-shell materialcould be identified to species. My studies of the cranial osteology of Galapagos tortoises (Crumly, 1984a) have notsupported Gunther's contention. Later, Giinther (1882) described Geoemyda [= Manouria] impressa from an 1 1 inchshell and noted its concave carapacial scutes. Gunther's outstanding illustrations of the type over-shadow theremainder of his description. Testudo kleinmanni was described by Lortet in 1883. His descrption emphasized shell color. Vaillant (1885)described Testudo [= Geochelone] yniphora on the basis of its not quite unique single gular scute (also found inChersina angulata and Geochelone vosmaeri). Vaillant also noted that the carapacial pattern was similar toGeochelone radiata. Femand Lataste described Homopus nogueyi [= Kinixys belliana nogueyi] in 1886. DespiteLataste's good description, he incorrectly assigned nogueyi to the genus Homopus. The small size of the typespecimens and the inadequate comparative material probably contributed to Lataste's error. Because of the smallnessof Lataste's specimens, they would not show the diagnotic feature, i.e., the carapacial hinge oi Kinixys.In 1886, Boulenger named three taxa all now included within Psammobates lentorius. Loveridge andWilliams (1957) considered Testudo fiski and Testudo smlthi synonyms of Psarnmobates tentorius verroxii, whereasBoulenger's T. trimeni was considered to be a valid subspecies oi Psammobates tentorius. Cervical scute size andshape, hip spine presence or absence, relative plastral scute length and shell color figured prominently inBoulenger's descriptions and good plates were included. Boulenger (1888) later described another species of smallSouth African tortoise, Homopus femoralis. Boulenger noted large hip spines, large prefrontal and frontal headscales, the absence of median lingual triturating ridge on the lower jaw, and five claws on the manus (unlike thefour-clawed condition in //. arelotus and//, signatus). In the following year, Baur (1889) discussed somenomenclatural problems in earlier works. As a consequence of his discussion, he proposed the names Testudo [=Geochelone] guentheri and T. [= Geochelone] galapagoensis. Unfortunately, nowhere did Baur discuss themorphological bases of his decisions. 1900-1950At the beginning of twentieth century, Rothschild named three new species from Galapagos in five shortnotes. In the first of these, Rothschild (1901) described Testudo [= Geochelone] becki on the basis of thecharacteristic flaring of the anterior carapace margin also found in other saddlebacked tortises. In his next two notes,Rothschild (1902a, 1902b) corrected some understandable mistakes concerning Harlan's (1827) type of Testudoelephantopus. In another paper, Rothschild (1902c) described Testudo wallacei, which has since been shown to be aninvalid, probably introduced, form (MacFarland et al. 1974a). Rothschild's final note (1903) concerned the tortoisespecies that now lives on Isla Santa Cruz (Indefatigable), which he named T. [= Geochelone] porteri (Fritts, pers.comm., believes this name is a junior synonym of T. [= Geochelone] nigrita).Friederick Siebenrock (1902) proposed the name Acinixys for Alfred Grandidier's Testudo planicauda andcompared it to Kinixys and the then unwieldly genus Testudo. Siebenrock (1903) later named Testudo [=Malacochersus] tornieri and noted its flattened and leathery shell, but considered it an abnormality. In the followingyear, Siebenrock's description (1904b) of Testudo boettgeri [= Psammobates tentorius verroxii] indicates hisconfusion with the astounding variation in some tortoise species. Also in 1903, Boulenger described Testudopseudemys [= Manouria impressa]. He used two features to distinguish T. pseudemys from similar species (M.emys and M. impressa). The length of the anterolateral vertebral sulci supposedly distingushed T. pseudemys fromM. impressa. However, this feature is quite variable and insufficiently delineated T. pseudemys from M. impressa.The narrow skull arches of T. pseudemys distinguished it from M. emys, which has broad skull arches. The skull ofT. pseudemys was unknown to Boulenger but has since been shown to be M. impressa.In 1906, Deurden named a new species of South African tortoise, Homopus boulengeri. Deurden noted thehigher number of marginal scutes (12 or 13, rarely 1 1), a small cervical scute, large imbricate forearm scales, andlarge hip spines, and several other features. Deurden also characterized the other species of Homopus, but wronglyincluded two taxa whose type specimens were juvenile Kinixys (H. darlingi Boulenger, 1902 and H. nogueyi Lataste,1886). Deurden (1907) more completely described the species of Homopus in the following year in a paperspecifically concerned with color pattern. Boulenger (1907) described Testudo [= Indotestudo] travancorica.Although he didn't make special note of the fact, he noted that the cervical scute was absent, unlike the condition inIndotestudo elongata. Boulenger (1907) included two excellent plates with his description; both showed thesimilarities with Indotestudo elongata. Boulenger made no mention of the earlier described species Testudo [=Indotestudo] forstenii Schlegel and Muller, 1840. Hoogmoed and Crumly (1984) showed that Indotestudo forsteniiwas the senior synonym of/, travancorica (Boulenger, 1907). Van Denburgh (1907) named four new tortoise speciesfrom Galapogos. All of his "diagnoses" were actually characterizations that relied mostly on shell ratios. However,both of the saddlebacked species described {Testudo [= Geochelone] phantasticus and Testudo [= Geochelone]hoodensis) were noted to have a throat marked with yellow. The throats of the other two species (7. [= Geochelone]darwinii and T. [= Geochelone] cliathamensis) were noted to be black. In 1913, Annandale, the superintendent of the Indian Museum, named Testudo [= Indotesiudo] parallelus fromChaibassa, Chota Nagpur. The specific name alludes to the parallel sides of the shell; other characters noted byAnnandale included gular, pectoral and humeral proportions. However, Annandale admitted that he was unable toobtain additional specimens besides the holotype. Testudo parallelus has been considered a synonym of/, elongata.During the same year that the British Museum shifted its collections to methyl alcohol and abandoned winespirits, George Boulenger continued his described two species of tortoises: Testudo buxtoni [= T. graeca ibera](1920a) and Testudo loveridgii [= Malacochersus tornieri] (1920b). Testudo buxtoni was distinguished from whatBoulenger considered its closest relatives (T. graeca ibera and T. graeca zarudnyi) by narrower pectoral scutes and adivided supracaudal scute; however, both the features are extremely variable in Testudo graeca. Testudo loveridgiiwas based on a series of six individuals; these individuals had divided supracaudal scutes. Arthur Lx)veridge (1923)reported on a collection of East African reptiles described and recognized Testudo procterae, differentiated fromBoulenger's Testudo loveridgii by its narrower, less flattened shell and its isolated distribution. Mertens et al. (1934)and Mertens and Wermuth (1955) recognized T. procterae as a distinct species and T. loveridgii a junior synonym ofM. tornieri. Both species were declared synonyms of Af. tornieri by Loveridge and Williams (1957).In 1929, Lindholm pubUshed a classification of turtles without supporting data. Therein, he recognized fivegenera (Testudo, Kinixys, Homopus, Pyxis and Acinixys) of tortoises and delineated four new subgenera of Testudo{Pampatestudo [= Chelonoidis], Indotestudo, Malacochersus and Goniochersus [= Chersina]). DeSola (1930) named aGalapagos species, Testudo [= Geochelone] vandenburghi, based on Van Denburgh's description (1914).In the early 1930's, John Hewitt described over a dozen new species and subspecies of tortoises, as well asthree new genera. Lx)veridge and Williams (1957) considered most of Hewitt's names to be junior synonyms of otherolder names. For example, Pseudomopus and Neotestudo were described in 1931 by Hewitt. Pseudomopus wasdistinguished from Homopus by its small inguinal scute and five toes of the manus. Neotestudo was proposed forChersina angulata, in order to remove this unique species from Testudo, at that time a large polyphyletic taxon.Hewitt (1931) also named two new subspecies oi Kinixys belliana and three new species oi Kinixys, all synonymsoi Kinixys belliana. In 1933, Hewitt named nine new subspecies of Chersinella [= Psammobates] tentorius. Eightof these subspecies are junior synonyms of Psammobates tentorius tentorius, fide Loveridge and Williams (1957).Hewitt (1933) proposed Megachersine [= Geochelone] for the species Geochelone pardalis and Geochelone elegansand Chersinella for the geometrica group of Testudo. Hewitts' generic descriptions relied on head scale variation.His new subspecies were based largely on small samples and used variable shell color and scale patterns.During his tenure as a Guggenheim Fellow, Loveridge collected a large number of reptiles and amphibiansfrom East Africa for the Museum of Comparative Zoology collections at Harvard University. His collectioncontained fourteen new taxa, including a new race of Geochelone pardalis. In 1935, Loveridge distinguished this racefrom the nominate form by its proportionally more vaulted shell and named it Testudo [= Geochelone] pardalisbabcocki. Grieg and Burdett (1976) suggested that the characters that distinguish the two subspecies of G. pardalisvary too much to warrant subspecific recognition.In a book on the fauna of Palestine, Friederich Bodenheimer described Testudofloweri [= T. graeca terrestris].His single sentence description, one of the shortest of the 20th century, is nevertheless sufficient: "The tiny varietyfound in the Negeb ..." (Bodenheimer, 1935). Hewitt described Kinixys natalensis in 1935 and noted that thesupracaudal scute is sometimes split. Loveridge and Williams (1957) considered K. natalensis d^i\m\ox synonym ofK. b. belliana. Broadley (1982) resurrected K. natalensis on the basis of pectoral scute proportions and a dividedsuprapygal. 1950-PresentIn his cervical vertebrae article Williams (1950a) suggested Bellemys as a substitute name for Pyxis. Asubsequent decision of the ICZN (Direction #32 of 1956) reversed Williams recommendation and Pyxis has beenretained. Heinz Wermuth, two years later (1952), named a new subspecies, Testudo hermanni robertmertensi in areview of T. hermanni from southern Europe. He distinguished this new subspecies on the basis of color pattern,particularly of the plastron, and overall shell proportions. In Laurent's (1956) survey of herpetofauna of the great lakes region of central Africa, he described a newsubspecies, Kinixys belliana mertensi. This report was soon followed by the most influential paper on Africantestudinids, Loveridge's and Williams' monograph on African cryptodires (1957). Their classification was a departurefrom earlier classifications (e.g., Mertens and Wermuth, 1955) which included most tortoises in the single largegenus Testudo. Instead, they recognized the so-called Ethiopian endemics {Chersina and Psammobates), elevatedGeochelone to generic level, and severely restricted the genus Testudo to include only north African, Mediterraneanand southwest Asian tortoises with a prootic concealed in dorsal view, a posterior plastral hinge, and "supranasal"scales. In addition, subgenera were proposed for Testudo kleinmanni (subgenus Pseudoiestudo) and Geochelonegigantea (subgenus Aldabrachelys). Loveridge disclaimed responsibility for these radical changes in classification ina footnote. He believed the changes would be overturned by future study; however, 1 have corroborated (Crumly,1984c) most of the changes suggested by Loveridge and Williams (1957).Wermuth (1958) usefully reviewed the nomenclature and status of Testudo graeca subspecies. He alsocorrected some nomenclatural errors and noted that Testudo terrestris ForskSl, 1775 (as a subspecies of T. graeca) isthe senior synonym oi Testudo flowed Bodenheimer 1935. Legler's (1959) decription of Gopherus flavomarginatusfollowed a year later. He distinguished it from other Gopherus on the basis of a longer intergular and a shorterinterhumeral sulcus, and a broad head. Khozatsky and Mlynarski (1966) suggested that Testudo horsfieldii beallocated to a new genus Agrionemys. They noted that T. horsfieldii had only four toes on the manus and that thedeeply "V" -shaped humeral pectoral sulcus laterally (but not medially) overlaps the entoplastron. My cladisticresults (Crumly, 1984b, 1984c) and Kirsch's (pers. comm.) breeding experiments do not support the recognition of aseparate genus for Testudo horsfieldii.In the late sixties and early seventies, Simone Vuillemin published several papers on Malagasian tortoises(1968 [with L. Rabodomihamina], 1972 [with C. Domergue], 1972a, and 1972b). Three new species were named inthe latter three papers. Unfortunately, Vuillemin apparently lacked sufficient comparative material, because all threeof his new taxa were clearly recognized as synomyms of earlier named species by both Obst (1978) and Bour (1979a,1981). Pyxoides brygooi Vuillemin & Domergue (1972) was considered a synomym of Pyxis arachnoides by Obstand was afforded subspecific rank {P. arachnoides brygooi) by Bour (1981), because it lacks an anteriorly mobilplastral lobe. Testudo morondavaensis Vuillemin (1972a), based on an immature specimen, was rightlysynonymized with Acinixys planicanda by both Bour and Obst. And Madakinixys domerguei Vuillemin (1972b),which was distinguished by dubious vertebral characters, was synonymized with Kinixys belliana by Obst (1978) andBour (1979a).Freiberg (1973) named two new species from southern South America, based on insufficient comparativematerial of Geochelone chilensis. The new species were named in honor of Dr. James A. Peters {Geochelone petersi)and Dr. Roberto Donoso-Barros {Geochelone donosobarrosi) and were distinguished by size, color, shape andthickness of the shell. G. petersi, a northern form of G. chilensis, has a smaller elongated, thick and light coloredshell; whereas G. donosobarrosi, a southern form of G. chilensis, has a large and dark-colored oval, but very thinshell. Marlow and Patton (1982) uncritically accepted Freiberg's names, but also showed that the three populationswere very closely related. My reasons for considering Freiberg's species as synonyms of Geochelone chilensis arediscussed in elsewhere (Crumly, 1984c). In short, Freiberg failed to note sexual dimorphism and latitudinal clinalvariation within G. chilensis, which could account for the variation he noted.The most active student of land tortoise evolution in the past few years has been Roger Bour of the MuseumNational d'Histoire Natureelle in Paris. His interest has been regional and limited to Madagascar and the islands ofthe Indian Ocean. Some of his papers have been reviews of the confusing nomenclatural history (1978, 1982b,1984a) of Malagasian testudinids. His most significant contribution has been to the systematics of Pyxis, anendemic tortoise genus of the Malagasy Republic (1979a, 1981, 1982a). In these papers, he recognized twosubspecies of Pyxis arachnoides and named a third that was later considered a synonym of an eariier name (Bour1982a). Thus, the trivial names recommended by Bour (1981, 1982a) are: Pyxis arachnoides arachnoides Bell, 1827;P. a. brygooi Vuillemin and Domergue, 1972; and P. a. oblonga Gray, 1869. In these same papers, Bourrecommended that Acinixys be considered a subgenus of Pyxis, citing P. a. brygooi as an intermediate betweenAcinixys and other Pyxis. For reasons presented elsewhere, I (Crumly, 1984c) prefer to retain Pyxis and Acinixys asseparate genera.Bour has also addressed the taxonomy of Indian Ocean tortoises in several papers. In 1982b, Bour rejected T.gigantea Schweigger, 1812, as the proper name for the tortoise species now inhabiting Aldabra and substituted T. elephantina Dum^ril and Bibron, 1835. He believed that this required the rejection of Aldabrachelys Loveridge andWilliams, 1957, and a new generic name. Thus, Bour coined Dipsochelys. However, there is an available genericname for tortoises with an elongated external narial opening: Megalochelys Falconer and Cautley, 1835 (type speciesM. atlas). I prefer to retain the specific epithet gigantea for three reasons: 1) an enormous amount of recentecological research has used the name, 2) abandonment of gigantea requires the acceptance of an uncertaininterpretation of Schweigger's (1812) description, and 3) elephantina, the next available name, is very much likeelephantopus and would increase the opportunity for orthographic errors (see Crumly, 1986). Still anotherinterpretation of Schweigger's original description is presented by Pritchard (1986); he suggests that Schweiggeractually described a specimen of Geochelone denticulata.Broadley (1982) reviewed the savannah Kinixys, some of which were subsumed into Kinixys helliana byLoveridge and Williams (1957). Broadley reinstated two names synonymized by Loveridge and Williams: Kinixysnatalensis (Hewitt, 1935) and Kinixys belliana spekii (Gray, 1863). K. natalensis was distinguished from otherKinixys by gular scute proportions, a tricupsid beak and a divided suprapygal. Kinixys belliana spekii wasdistinguished from other K. belliana by carapacial and gular proportions, a unicuspid beak and coloration.Bramble (1982) restricted Gopherus by removing two of the primitive species and placing them in his newgenus Scaptochelys. Although Gopherus (sensu lato) is exceptionally well defined, Scaptochelys is based on sharedprimitive characters (Crumly, 1984b, 1984c). Furthermore, Scaptochelys, containing Gopherus agassizii and G.berlandieri as well as some fossil species, is a junior subjective synonym of Xerobates Agassiz 1857 (type speciesXerobates berlandieri designated by Brown 1908) (Bour and Dubois, 1984; Crumly, 1985).Selected LiteratureBelow are the references cited in the preceeding pages. Also included are other references that were not cited,but that are useful for the interpretation of the history of testudinid nomenclature. This is not intended as a completebibliography of land tortoise systematics. Those references cited in the text are preceded by an asterisk.Agassiz, L. 1857. Contribution to the Natural History of the United States. North American Testudinata. 1 (2):235-452. Little, Brown and Co.; Boston.Anderson, J. 1898. Zoology of Egypt, 1, Replilia and Balrachia. Bernard Quartich; London.Anderson, S. C. 1966. The turtles, lizards and amphibaenians of Iran. Ph.D. Dissertation, Stanford University.Andrews, C. 1906. A descriptive catalogue of the Tertiary Vertebrata of the Fayum, Egypt. British Mas. (Natur.Hist.); London.Annandale, N. 1906. Testudo baluchiorum, a new species. J. Proc. Asiatic Soc. Bengal 2(3): 1-11. . 1913. The tortoises of Chola Nagpur. Rec. Indian Mus. 9(2), no. 5: 63-78. . 1915. Notes on some Indian chelonians. Rec. Indian Mus. part 11 11(11): 189-195.Arnold, E. N. 1979. Indian Ocean giant tortoises: their systematics and island adaptations. Phil. Trans. R. Soc.London B. 286: 127-145.Auffenberg, W. 1961. A correction regarding the phalangeal formula of the turtle Stylemys nebrascensis Leidy.Copeia 1961(4): 496-498. . 1962. A new species of Geochelone from the Pleistocene of Texas. Copeia 1962(3): 627-636. . 1963. The fossil testudinine turtles of Florida, genera Geochelone and Floridemys. Bull. Florida StateMus. 7(2): 53-97. . 1964. A new fossil tortoise from the Texas Miocene, with remarks on the probable geological historyof tortoises in the Eastern United States. The Pearse-Sellards Series, Texas Mem. Mus., Univ. Texas3: 2-10. . 1966. The carpus of land tortoises (Testudininae). Bull. Florida State Mus. 10(5): 159-191. . 1969. Land of the Choco tortoise, Geochelone chilensis. Intematl. Turtle & Tortoise Soc. J. 3(3). . 1971. A new fossil tortoise, with remarks on the origin of South American tortoises. Copeia 1971(1):106-117.* . 1974. Checklist of fossil land tortoises (Testudinidae). Bull. Florida State Mus. 18(3): 121-251. __. 1976. The genus Gopherus (Testudinidae): Pt. I. Osteology and relationships of extant species. Bull.Florida State Mus. 20(2): 47-110.and R. Franz. 1978a. Gopherus. Catalogue Amer. Amphibians Reptiles (211): 1-2.and . 1978b. Gopherus agassizii. Catalogue Amer. Amphibians Reptiles (212): 1-2.and . 1978c. Gopherus berlandieri. Catalogue Amer. Amphibians Reptiles (213): 1-2.and . 1978d. Gopherus flavomarginatus. Catalogue Amer. Amphibians Reptiles (214): 1-2. 1978e. Gopherus polyphemus. Catalogue Amer. Amphibians Reptiles (215): 1-2.Bartram, J. 1791. Travels through North and South Carolina, Georgia, East and West Florida ... Containing anAccount of the Soil and Natural Productions of Those Regions. James and Johnson; Philadelphia[1955 Dover Publications Inc. reprint].+Baur, G. 1889 (1890). The gigantic land tortoises of the Galapagos Islands. Amer. Naturalist 23(276): 1039-1057. . 1892. Bermerkungen iibcr verschiedene Arten von Schildkroten. Zool. Anz. (389): 1-5.*Bell, T. 1827. On two genera of land tortoises. Trans. Linn. Soc. London 15(2): 392-401.* . 1828. Descriptions of three new species of land tortoises. Zool. J. London 3: 419-421.1836. A Monograph of the Testudinata. Samuel Highley; London.Berthold, A. A. 1845. Uber Cinixys homeana Bell. Nova Acta Acad, leop.-carol. 22: 423-428.Blanford, W. T. 1876. The zoology and geology. Pp. 305-436 In: F. J. Goldsmid (ed.), Eastern Persia: An Accountof the Journeys of the Persian Boundary Commission 1870-71-72. vol. 2. London; Oxford.*Blyth, E. 1853. A collection of reptiles from Afghanistan. J. Asiatic Soc. Bengal 22: 640. . 1854. Report of the curator (Zoological Dept., Parts: Reptiles and Amphibians). Proc. Asiatic Soc.Bengal 23: 737-740.* . 1863. Notes on Asiatic reptiles. J. Asiatic Soc. Bengal 32: 83.*Bodenheimer, F. S. 1935. Animal Life in Palestine. L. Mayer; Jerusalem.Boettger, O. 1893. Katalog der Reptilien-Sammlung im Museum der Senckenbergischen Naturforschenden Gesellschaftin Frankfurt am Main. 1. Rhynchocephalen, Schildkroten, Eidechsen iind Chamaleons. GebriiderKnauer; Frankfurt am Main.Bonnatcrre, A. 1789. Tableau Encyclopedique et Methodique des Trois Regnes de la Nature. Panckoucke; Paris.*BouIenger, G. A. 1886. On the South African tortoises allied to Testudo geometrica. Proc. Zool. Soc, London:540-542.* . 1888. Description of a new land-tortoise from South Africa, from a specimen living in the Society'sgarden. Homopus femoralis. Proc. Zool. Soc. London: 251.* . 1889. Catalogue of the Chelonians, Rhynchoccphalians, Crocodilians in the Collection of the BritishMuseum (Natural History). Trustees of the British Museum, London. . 1902. (Description of Homopus darlingi). Proc. Zool. Soc. London (2): 15. . 1903a. Report on the batrachians and reptiles. Pp. 131-176 In: N. Annandale and H. C. Robinson(eds.). Fascic. Malay. Zool., 1. . 1903b. On a collection of batrachians and reptiles from the interior of the Cape Colony. Ann. Mag.natur. Hist. London (7), 12: 215-217.* . 1907. A new tortoise from Travancore. J. Bombay Natur. Hist. Soc. 17: 560-561. . 1920a. Description of a new land tortoise from northern Persia. Proc. Bombay Natur. Hist. Soc.27(2): 251-252.* . 1920b. Une Tortue extraordinaire: Testudo loveridgii, sp. n. C. R. Acad. Sci. (Paris) 170: 263-266.Bour, R. 1978. Les tortues des Mascareignes; description d'une espfece nouvelle d'apr^s un document (Mdmoires deI'Acaddmie) de 1737 dans lequel le crane est figure. C. R. Acad. Sc. Paris 287: 491-493.* . 1979. Les tortues actuelles de Madagascar (Republique Malgache): Liste systSmatique et description dedeux sous-esp6ces nouvelles (Reptilia-Testudines). Bull. Soc. Et. sci. Anjou, N.S. 10: 141-154. . 1979b. Premiere decouverte de restes osseux de la Tortue terrestre de la Reunion, Cylindraspisborbonica. C. R. Acad. Sc. Paris 288: 1223-1226. . 1980. Essai sur la taxinomie des Testudinidae actuels (Reptilia, Chelonii). Bull. Mus. nail. Hist, natur.Paris 4, 2, A(2): 541-546. . 1981. Etude syst6matique du genre enddmique Malagache Pyxis Bell, 1827 (Reptilia, Chelonii). Bull.Soc. Linn. Lyon 50(4): 132-176. . 1982a. Etude systematique du genre endfimique Malagache Pyxis Bell, 1827 (Reptilia, Chelonii). (Notecomplementaire). Bull. Soc. Linn. Lyon 51(1): 28-31. . 1982b. Contribution & la connaissance des Tortues terrestres des Seychelles: definition du genereendSmique et description d'une espfece nouvelle probablement originaire des iles granitiques et au bordde I'extinction. C. R. Acad. Sc. Paris 295, ser. Ill, 121-122.* . 1984a. Taxonomy, history and geography of Seychellois Land Tortoises and fresh water turtles. Pp.281-307 In: D. R. Stoddart (ed.), Biogeography and ecology of the Seychelles Islands. Dr. W. JunkPubl.; The Hague.* . 1984b. L'identite de Testudo gigantea Schweigger, 1812 (Reptilia, Chelonii). Bull. Mus. natl. Hist.natur. Paris 4, 6, A(l): 159-175.* and A. Dubois. 1984. Xerobates Agassiz, 1857, synonyme plus ancien de Scaptochelys Bramble, 1982(Reptilia, Chelonii, Testudinidae). Bull. Soc. Linn. Lyon 53(1): 30-32.Bourret, R. 1941. Les Tortues de L'Indochina. Inst. Oceanographique L'Indochina (38): 1-235.Bramble, D. M. 1971. Functional morphology, evolution and paleontology of Gopher tortoises. Ph.D. Dissertation,Univ. of California, Berkeley. 1974. Emydid shell kinesis: Biomechanics and evolution. Copeia 1974(3): 101-127.* . 1982. Scaptochelys: generic revision and evolution of Gopher tortoises. Copeia 1982(4): 853-866.Brattstrom, B. H. 1961. Some new fossil tortoises from western North America, with remarks on the zoogeographyand paleoecology of tortoises. J. Paleo. 35(3): 543-560.*Broadley, D. G. 1982. A review of the populations of Kinixys (Testudinidae) occurring in south-eastern Africa.Ann. Cape Prov. Mus. (Natur. Hist.) 13(14): 195-215.Broin, F. de. 1977. Contribucion i L'fitude des Ch^loniens: ch^loniens continentaux du CrStac6 et du Tertiaire deFrance. M6m. Mus. natl. d'Hist. natur., ser. C, 1-366.Carr, A. 1952. Handbook of Turtles: Turtles of the United States, Canada and Baja California. Cornell UniversityPress, Ithaca.Chenu, J. C. 1857. Encyclopedie D'Histoire Naturelle. Reptiles et Poissons. Avec la collaboration de M. E.Desmarest. Marescq; Paris.Cheylan, M. 1981. Biologic et ecologie de la tortue d'Hermann Testudo hermanni, Gmelin 1789. Contribution deI'espfece a la connaissance des climats Quatemaires de la France. M6m. Trav. E. P. H. E., Inst.Montpellier 13: 1-404.Chkikvadze, V. M. 1970. [On the origin of modem palaearctic land tortoises]. Bull. Acad. Sci. Georgian S. S. R. 57(1): 245-247 [in Russian, English abstract]. . 1972. [On the systematic position of the Tertiary gigantic land tortoises of the Palaearctic]. Bull.Acad. Sci. Georgian S. S. R. 65 (3): 745-748 [in Russian].Cochran, D. M. 1961. Type specimens of reptiles and amphibians in the United States National Museum. Bull.United States Natl. Mus. (220): 1-291.1876. [A letter from Cookson to A. GUnther read to the Society]. Proc. Zool. Soc. London (35):520-526.1863. New California animals. Proc. California Acad. Sci. 2: 118-123.1980. The cranial osteology and evolution of the tortoise genus Geochelone (Testudines,Testudinidae). M.S. Thesis, San Diego State University.1982a. A cladistic analysis of Geochelone using cranial osteology. J. Herpetol. 16(3): 215-234. _. 1983. An annotated checklist of the fossil tortoises of China and Mongolian. Proc. Biol. Soc.Washington 96(3): 567-580. _. 1984a. The cranial morphometry of Galapagos tortoises. Proc. California Acad. Sci. 43(9): 111-121. _. 1984b. A hypothesis for the intergeneric relationships of land tortoises (family Testudinidae). FirstIntematl. Syrap. Fossil Turtles, Studia Geologica, Salmanca, vol. esp. 1: 115-124. _. 1984c. The evolution of land tortoises (family Testudinidae). Ph.D. Dissertation, Rutgers - The StateUniversity. _. 1985. The genus name for North American Gopher tortoises. Proc. Desert Tortoise Council 1984Symp.: 147-148. _. 1986. The identy of Testudo gigantea Schweigger, 1812: Another interpretation. Herpetologica42(2): 237-241.1829. La Rfegne Animal distribufe d'aprSs son Organisation, pour servir de base a I'Histoire naturelle deAnimaux et d'Introduction k 1'Anatomic comparee. ed. 2, Paris, Deterville, 2, Les Reptiles. Cookson, W, E.Cooper, J. G.Crumly, C. R. *Darevsky, I. S. and R. Mertens. 1973. Zwei unveroffentlichte Schildkrotentafeln von Pallas. Salamandra 9(3/4): 99-102.Daudin, F. M. 1802. Histoire naturelle, generate et particulaire des reptiles, vol. 2: 1-432. F. Dufart; Paris.Deraniyagala, P. E. P. 1939. The Tetrapod Reptiles of Ceylon, vol. I. Testudinates and Crocodilians. Ceylon J. Sci.Colombo Mus. Natur. Hist. 1-412. . 1953. A Colored Atlas of Some Vertebrates from Ceylon. II. Tetrapod Reptilia. Natl. Mus. Ceylon;Colombo.DeSola, R. 1930. The Liebspiel of Testudo vandenburghi, a new name for the mid-Albemarle Island Galapagostortoise. Copeia 1930(3): 79-80.DcWaal, S. W. P. 1981. The Testudines (Reptilia) of the Orange Free State, South Africa. Navorsinge Nasionale Mus.Bloemfontein 4(3): 85-91.*Duerden, J. E. 1906. South African tortoises of the genus Homopus, with descriptions of a new species. Rec.Albany Mus. 1: 405-411.* . 1907. Variations in the Geomeirica group of South African tortoises. Rep. S. African Assoc. Advanc.Sci., 1905-1906: 178-208.*Dum6ril, A. M. C. and G. Bibron. 1835. Erpetologie generale ou histoire naturelle complete des reptiles, vol 2.Librairie Encyclop^dique du Roret; Paris.Dum^ril, A. M. C, G. Bibron, and A. H. A. Dum6ril. 1851. Catalogue methodique de la collection des reptiles duMuseum d'Histoire Naturelle de Paris. Gide et Baudry; Paris. Edwards, G. 1751. A Natural History of Birds. Part 4. 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Notes on a collection of reptiles and amphibians from the west coast of southern Africa. Ann.Transvaal Mus. 21: 253-259.Flower, S. S. 1925. Contributions to our knowledge of the duration of life in vertebrate animals. III. Reptiles. Proc.Zool. Soc. London: 911-981. . 1933. Notes on the recent reptiles and amphibians of Egypt, with a list of species recorded from thatkingdom. Proc. Zool. Soc. London: 735-851.Forskal, P. 1775. Descriptiones Animalium, Avium, Amphibiorum, Piscium, ... post mortem auctoris editit C.Nieburh. 4 vols. Hauniae; Copenhagen.Freiberg, M. 1973. Dos nuevas tortugas terrestres de Argentina. Bol. Soc. Biol. Concepcion 46: 81-93.Fritts, T. H. 1983. Morphometries of Galapagos tortoises: evolutionary implications. Pp. 107-122 In: R. I.Bowman, M. Berson & A. E. Leviton (eds.), Patterns of Evolution in Galapagos Organisms, PacificDiv., Amer. Assoc. Advanc. Sci., San Franciso.Carman, S. 1917. The Galapagos tortoises. Mem. Mus. Comp. Zool. 30: 262-296.Gilmore, C. W. 1915. Fossil turtles from the Uinta formation. Mem. Carnegie Mus. 7 (2): 101-161. . 1931. Fossil turtles of Mongolia. Bull. Amer. Mus. Natur. Hist. 59: 213-257.Gmelin, J. F. 1789. In: C. Linnaeus, Systema Naturae. 13th edition, 1-1053. . 1831. In: J. E. Gray, Synopsis Reptilium. London. 1-85. . 1839. Naturgeschichte der Amphibien. Mannheim. 1-22.Grandidier, A. 1867. Liste des reptiles nouveaux discouverts en 1866 sur la cote Sud-Ouest de Madagascar. Rev.Mag. Zool. July 1867: 232-234.Gray, J. E. 1828, 1830. Spicilcgia Zoologica; or Original figures and short systematic descriptions of new andunfigured animals. Part I (1828), Part II (1830), 1-12. [Plates i-ix were never published, but are at theBritish Museum]. Trustees of the British Museum; London. . 1831a. Synopsis of the species of the Class Reptilia. Pp. 1-481 + l-IIO, pis. i-x!ii. In: E. Griffith,The Animal Kingdom arranged in conformity with its organization by Baron Cuvier. Whittaker,Treacher and Co.: London. . 1831b. 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