We note that despite the extensive discussions and opinions on this issue, one rare
point of general agreement seems to be to accept that there is only a single species of
‘giant land tortoise’on Aldabra Island, so the problem simply amounts to knowing
what it should be called. If we include the original application by Frazier, in 2009 no
fewer than 91 persons commented so far on this case in 72 contributions in four
instalments of this Bulletin, covering 66 pages. We note that 85 of the intervening
parties were in favour of the name Testudo gigantea for this species and 6 were in
favour of the name Testudo dussumieri. However, we are convinced that nomenclatural
decisions by the Commission should not rely on polls or on persons of variable
expertise and insight, nor on campaigns seeking to gather supporters to form a
‘pressure group’, but should be based upon due consideration of explicit arguments,
even if expressed by a ‘minority’ of stakeholders.
As the present curator (AO) and the previous two curators (AD, ERB) of the
herpetological collection of the Muséum National d’Histoire Naturelle (Paris), one of
the oldest and most important herpetological collections in the world, we agree with
Gerlach’s statement that the comments in favour of each name for the Aldabra
tortoise rest on (1) taxonomic arguments regarding the identifications of the type
specimens; (2) arguments for ‘nomenclatural stability’, relating to usage of names,
irrespective of the identification of the taxa; (3) arguments concerned with the
conservation biology of these tortoises and (4) arguments relating to the appropriateness
of each name for the taxon. Let us consider these four sets of arguments in the
Gerlach and Hansen discussed the respective appropriateness of the names gigantea
and dussumieri, and Aldabrachelys and Dipsochelys. This discussion is totally
irrelevant to the present issue. As pointed out by Dubois & Raffaëlli (2009, p. 18),
scientific names are not descriptions, diagnoses, statements of characters, distribution
or other characterisations of the taxa they designate, nor models or theories about the
hypothesised origin of these taxa, nor praise for their authors, for the discoverers of
the taxa or for the persons to whom they may be dedicated. They are just neutral
labels designating unambiguously and universally given taxa within the frame of a
given taxonomy, i.e. allowing automatic reference to the taxa recognised by
taxonomists at a given stage of their research. These labels allow storage and retrieval
of the information accumulated in taxonomies, but it is not mandatory that the
names have any meaning at all. In many cases, factually true information about
coloration, body form or geographic range may indeed be encapsulated in either or
both of the two terms of a binomen, but in many other cases the name provides
misleading statements about the taxon. This is why the Code expressly states that
availability (and consequently validity) of names ‘is not affected by inappropriateness’
(Article 18), and allows a new generic or specific name to be ‘empty of meaning’,
for example for being ‘an arbitrary combination of letters provided this is formed tobe used as a word’ (Article 11.3). If it were not so, thousands of zoological names
would have to be changed. There is no point in further discussing the question of
appropriateness of names in the present case.
This was referred to by 42 of the 85 (49.4 %) supporters of the application, but, as
correctly stated by Gerlach (BZN 66: 184–186, June 2009), it is not convincing: ‘the
tortoises referred to are explicitly the Aldabra tortoises, for which there is no
significant identification issue whatever name is applied’. Whenever a decision is
made regarding the valid name of this species, it will be a trivial matter to incorporate
it in official checklists and documents, as rightly stressed by Hoogmoed (BZN 66:
354–356): ‘international bodies like CITES are able to change names of species on
their lists with few problems and without jeopardising the protection of the taxa in
question. And the same holds true for governments and their agencies’. Such changes
of names of taxa on official lists have already occurred on several occasions, without
causing any problem for the conservation policy of the taxa concerned. As long as
the species is well identified, a unique and universal name is not essential for the
conservation of threatened taxa. Whereas the Aldabra tortoise is in Annex 2 of the
CITES list, the following three taxa are in Annex 1 and their names, long considered
to be ‘universal’, have changed during the last 50 years: the mammals Papio sphinx
(Linnaeus, 1758) and Gazella dama (Pallas, 1766) have become respectively Mandrillus
sphinx and Nanger dama; the bird Diomedea albatrus Pallas, 1769 changed into
Phoebastria albatrus; the fish Pangasionodon gigas Chevey, 1931 is now known as
Pangasius gigas; and the chelonians Kachuga tecta (Gray, 1831) and Testudo
elephantopus Harlan, 1827 are now on the list respectively as Pangshura tecta and
Chelonoidis nigra (Quoy & Gaimard, 1824). The last example is very similar to that
of the Aldabra tortoise: it is also a threatened insular giant tortoise of which both the
generic and specific names were changed. In some other cases a single species appears
under different names on different lists: for example, the bird mentioned on the
CITES list as Houbaropsis bengalensis (Gmelin, 1789) appears on the IUCN list as
Eupodotis bengalensis. We know of no evidence that these changes caused any
problem for the conservation policy of these species.
Of course, we agree with Vences (BZN 66: 282, September 2009) that, for
conservation (but also other) purposes, stabilising the nomenclature of the Aldabra
tortoise will ultimately be important, and that, now that Frazier has challenged the
correct nomenclature, it is unlikely that the scientific community will reach a
consensus on which name to use without an unambiguous decision of the Commission.
But this does not imply in the least that this decision should follow Frazier’s
suggestion. The merits of Frazier’s proposal have to be evaluated in the light of the
next two arguments.
Among the 85 persons who expressed their support for the use of the name T.
gigantea, 59 (69.4 %) did not challenge the respective identifications of the lectotype
of T. dussumieri presented by Bour (1984) or of the holotype of T. gigantea presented
by Pritchard (1986) and Bour (2006). Their opinion rested on the assumption thatusage of T. gigantea is sufficiently well established to require conservation of this
name through use of the plenary power of the Commission, whatever species its
holotype represents. However, comments by Bour & Pritchard and Cheke (BZN 66:
174–176, June 2009) disagreed with this assumption. The statement by Frazier
regarding usage of the name gigantea was clearly demonstrated to be in error by
Bour, Pritchard & Iverson (BZN 67: 73–77, March 2010), through a survey of
Google. In recent years, this single species has been designated mostly under 3
different specific names, dussumieri, elephantina and gigantea, and 3 different generic
names, Aldabrachelys, Dipsochelys and Geochelone. These data show: (1) that no
universality of usage exists regarding these names; (2) that the relative numbers of
authors using each of these names are constantly changing; and (3) that, during
recent months, the relative frequency of usage has increased for gigantea and
Aldabrachelys, but decreased for dussumieri and Dipsochelys. As a matter of fact, the
figures obtained about nine months after publication of Frazier’s application show a
strong impact of this application itself upon usage. In addition to the content of the
application itself and personal contacts of its author with colleagues, this is clearly
due to (1) the fact that the title of this application contains the misleading statement
‘currently Geochelone (Aldabrachelys) gigantea’, and (2) the fact that the Code states
that while such a case is under consideration the invalid name (under the normal
rules) has to be used! On the last check-list of the extant turtles and tortoises by
Rhodin et al. (2009), contrary to that by Fritz & Havas (2007), both combinations
Aldabrachelys gigantea and Dipsochelys dussumieri are proposed together as an
alternative to name the Aldabra tortoise. Pending the decision of the Commission,
this appears to us the best attitude to adopt in the present highly controversial
The arguments of the supporters of Frazier’s application are mostly directed
against the name dussumieri, but this is a biased presentation of the facts. The name
dussumieri was resurrected as the valid one for the Aldabra tortoise only in 1995
(Gerlach & Canning, 1995; Gerlach, 1997), but the fact that the name gigantea does
not apply to the Aldabra tortoise had been established 13 years earlier, by Bour
(1982). Bour had made an error concerning the biological species to which the
holotype of gigantea belonged, an error corrected by Pritchard (1986) twenty years
before the rediscovery of the holotype specimen by Bour (2006), but nevertheless it
has been clear from 1982 that the name gigantea, created for a tortoise from Brazil,
does not apply to the Aldabra tortoise. Therefore, between 1982 and 1995, pending
the resurrection of the name dussumieri, it was normal and correct to use the name
elephantina for this species, which explains why 18.5 % of the Google hits obtained
by Bour, Pritchard & Iverson (BZN 67: 73–77, March 2010) concern this name. The
Seychelles Island Foundation (SIF), some twenty members of which signed comments
in support of the use of Geochelone gigantea, still recently used official
documents where the Aldabra tortoise was named either Dipsochelys dussumieri or
Testudo elephantina (e.g. Beaver & Gerlach, 1998; Anonymous, 2001). Furthermore,
and contrary to the statements of Frazier and his supporters, the name elephantina
had also been used a long time prior to 1982 by some authors, and there was no
period in history when the name gigantea was the only one used for the Aldabra
tortoise. During the so-called period of ‘universality’ of usage of the name gigantea,
the name elephantina was regularly used as valid for a species or subspecies by a
84 Bulletin of Zoological Nomenclature 67(1) March 2010
minority of authors: e.g. Fritsch, 1871; Günther, 1877; Peters, 1882; G.A. Boulenger,
1889; Strauch, 1890; Schacht, 1903 (as Testudo elefantina); E.G. Boulenger, 1914;
Geiman & Wichterman, 1937; Evans & Quaranta, 1949; Quaranta & Evans, 1949;
Urbain et al., 1951; Wermuth & Mertens, 1961; Georg et al., 1962; Steers, 1968.
The Glossary of the Code (p. 121) defines ‘prevailing usage’ of a name as follows:
‘that usage of the name which is adopted by at least a substantial majority of the most
recent authors concerned with the relevant taxon, irrespective of how long ago their
work was published’. This definition is not clear, as it does not provide guidelines to
recognise ‘the most recent authors’ and ‘at least a substantial majority’, but the
minimum that the latter words may mean is that this is a majority of ‘much more
than 50 %’. It is therefore clear that the argument of ‘usage’ does not hold in the
present case. There has never existed any ‘universality of usage’ for the scientific
name of the Aldabra tortoise, but a diversity of usages, and stabilisation of usage in
this case should come from simply following the Code. If ‘usage’ of a specific name
in 46.3 % of recent publications and internet documents, including many posterior to
an application urging the Commission to stabilise this usage, against 35.3 and 18.5 %
for two other names (see Bour, Pritchard & Iverson, 2010), could be retained to
nullify the rules of zoological nomenclature, then we might as well get rid of these
rules altogether, including the Principle of Priority and, why not, the entire Code
itself, leaving so-called ‘consensus’ to decide upon the valid names of taxa.
Identification of the holotypes
Nineteen (22.4 %) of the 85 supporters of the name T. gigantea challenged the
identification of the holotype of T. gigantea and one (1.2 %) did so for the lectotype
of T. dussumieri. Because giant insular tortoises tend to share many homoplasies due
to similar environmental conditions, we consider that the specific identification of
museum specimens can be relied upon only if carried out by experienced taxonomists
well-acquainted with the group of modern land tortoises. This is stressed by a droll
example: one month after publication in this Bulletin of his support for the original
application regarding this case, a museum curator sent the Paris Museum for
identification a set of photos of a specimen of giant land tortoise, stating that he was
‘not too sure’ about what this specimen could be. Well, this specimen happened to be
unquestionably . . . an Aldabra tortoise!
Among the 91 persons who commented on this case, only three have personally
examined the holotypes of the two nominal species here at stake. The lectotype
RMNH 3231 of Testudo dussumieri Gray, 1831 has been examined only by
Hoogmoed (then curator of the RMNH collection), Bour (1984) and Pritchard
(during a visit to the Leiden Museum in 2000), and the holotype MNHN 9566 of
Testudo gigantea Schweigger, 1812 has been examined so far only by Bour (2006),
although Pritchard has examined an extensive set of colour photographs of this
specimen, sent to him by Bour. All other authors who commented on the taxonomic
allocation of these specimens did so only on the basis of the original descriptions
(which they presumably read) or possibly of published photographs of the holotypes,
if they indeed had access to their publication in a little known journal (Bour, 2006).
None of them ever wrote to our Museum to request access to the holotype of
T. gigantea, or to photographs, radiographs or measurements of it, and we note with
some regret that perhaps even the assumption that the contributors to this discussion
Bulletin of Zoological Nomenclature 67(1) March 2010 85
read Schweigger’s original work, as well as those of Pritchard and Bour, may be in
error for some of them.
Among the 16 testudinid taxonomists involved in the discussion beside Bour, 5
expressly mentioned that they agreed with Bour’s (2006) statement that the specimen
MNHN 9566 is indeed the specimen that had been described by Schweigger (1812),
6 did not question this statement, and 5 (Arnold, Fritz, Meylan, Parham & Rhodin)
expressed scepticism about it, but did not identify a single morphological or
anatomical character of this specimen that was not compatible with the original
description. Another taxonomist (Matyot) questioned the origin of the holotype of
T. dussumieri, but did not provide a reasonable argument to back up this opinion (see
Bour, Iverson & Pritchard, 2010).
Finally, none of these supporters of Frazier’s application provided any argument
to explain the discrepancies between the original description of Testudo gigantea by
Schweigger (1812) and the biological species of the Aldabra tortoise. Therefore,
following the precise conditions put by Article 75.3 of the Code for allowing a
neotype designation, it is clear that, as stated by Bour & Pritchard, Cheke, Gerlach
and Hoogmoed, the designation of a neotype for this species by Frazier (2006) is
invalid, as this specimen does not come from the original type locality (Brazil) and
differs in several important characters (see the comments by Bour & Pritchard and by
Gerlach) from the original description. Nobody in the world can agree that a
specimen from Aldabra qualifies as coming ‘as nearly as practicable from the original
type locality’ (Article 75.3.6) of a tortoise from Brazil. Therefore, even if the holotype
of this nominal species had not been rediscovered, this designation would be null and
void, and should have been replaced by another one based on a tortoise specimen
from Brazil or, if this had turned impracticable (e.g. because of restrictive laws on the
export of specimens of this endangered species), from a neighbouring country.
Therefore, we concur with Bour & Pritchard, Cheke, Gerlach, Hoogmoed and
Iverson that no evidence has been provided by the supporters of the application that
Bour’s (1984, 2006) taxonomic interpretations of the holotype of T. gigantea and of
the lectotype of T. dussumieri are incorrect. These data dictate that the Commission
should not use its plenary power to set aside all previous type fixations, validate
Frazier’s neotype for the former, and suppress the latter.
As present or past curators of an important natural history museum collection, we are
quite worried about the turn that this discussion has taken. Most of its actors based
their comments on opinions, tastes, or just ‘deep feelings’, rather than on the respective
merits of rational arguments. In particular, we were very shocked to see the publication
of personal attacks against our colleague Roger Bour, questioning his honesty and
suggesting that he manipulated scientific data. These published statements will remain
available in the literature long after the death of all contributors to this discussion. We
have known Roger for about 40 years and we would like to praise his scientific
competence, his intellectual honesty and his refusal to make ‘political calculations’ to
allow his opinions to win or to enhance his career. He has devoted a lot of his
professional life to clarifying difficult taxonomic and nomenclatural problems in chelonians,
identifying old ‘forgotten’ types in many museums worldwide, and thus permitting
genuine nomenclatural stabilisation based on scientific data, not on ‘impressions’,
86 Bulletin of Zoological Nomenclature 67(1) March 2010
‘profound hunches’ or ‘lobbies’. We understand these repeated attacks as a disapproval
and a denial of the quality of his work, and we think he deserves better treatment from
the part of the many colleagues worldwide who benefited from his help in their research
for decades. We consider this failure as a pernicious result of the opening of this
unwarranted case in this Bulletin, which is itself a consequence of the emphasis put by
the Commission in recent years on ‘usage’ against the rules of the Code.
We agree with Hoogmoed’s statement that ‘Frazier’s proposal is completely
unnecessary, because the facts are clear and the rules of the [Code] provide solutions
for this situation’. The case at stake here in fact concerns neither conservation biology
nor nomenclatural stability, but simply the accuracy of taxonomic work.
The doubts cast by some authors about the rediscovery of a holotype in an old
collection like that of the Paris Museum demonstrate a poor knowledge about such
historical collections. As experienced taxonomists, we had on various occasions the
opportunity to rediscover specimens that were not labelled as types, not only in the
Paris Museum but also in other old historically important collections, like those of
London or Berlin. An important part of the herpetological collection of the Paris
Museum, but even more so of the mammal and bird collections of that institution, is
composed of historical specimens, many of which can potentially be name-bearing
types, or at least vouchers of specimens mentioned in ancient publications. Such
publications date from the end of the 18th and the 19th century, when no such
regulations as the Code existed and no proper labelling of specimens as ‘types’ could
be done, as such a concept did not exist or was used in a very vague way (e.g. an
author could then decide to replace the original ‘type’ by a ‘more appropriate’
specimen). In a work in progress, the type catalogue of hyloid frogs in the Paris
Museum herpetological collection (Ohler et al., in preparation) covers 156 names; 67
(43 %) were created in works published before 1854, and 90 (58 %) before 1900, at the
time of implementation of the first Code (the ‘Règles’). A similar proportion of old
names would probably be found in other parts of the collection, or this proportion
may even be higher for a well-studied group like the chelonians. Some of the
intervenors in this debate seemed to consider that it would be a crime of lese-majesty
to consider that Duméril & Bibron, ‘these two doyens of herpetology’ (Lenin &
Frazier, 2009) could have made an identification error on a specimen. However,
everybody can make a mistake and, considering the monumental work they
produced, there is nothing surprising or shocking to note that Duméril & Bibron
(1834–1854) made a number of mistakes, not only about the identity of the holotype
of Testudo gigantea (e.g. Shea, 2001; Lescure & Ohler, in preparation).
Old museum collections covering the whole of zoology still harbour thousands of
historical specimens, including name-bearing types that have not yet been identified as
such. Every thorough survey of old specimens of any zoological group in such museums
is an adventure which can be as exciting as field work in the remotest places of the earth.
Doing so, one sometimes finds unexpected results, e.g. regarding the taxonomic identification
of old name-bearing types, and some names have to change. The increased
availability and application of DNA sequencing technology to the proper taxonomic
identification of name-bearing types promises that the frequency of nomenclatural
complications will only increase. Should curators and taxonomists stop exploring these
resources to avoid such unexpected findings? Should they throw away these old
specimens for fear that they would upset ‘usage’ of the names at stake and then threaten
Bulletin of Zoological Nomenclature 67(1) March 2010 87
the personal comfort of some persons long involved in researches dealing with these
animals? Should we just close museums in order to please ‘conservationists’ (of names)?
Bouchet (BZN 66: 77, March 2009) rightly stressed that ‘Vertebrate paleontology
survived the name Brontosaurus giving way to Apatosaurus’, and it can be quite safely
added that the extinction of these animals was not caused by this synonymisation. All
palaeontologists now use the name Apatosaurus for this genus, and this change did not
create problems for non-taxonomists, who may still use the common English name
‘brontosaurs’ for these animals. Whatever scientific name will ultimately be retained for
this species, the Aldabra tortoise is and will remain designated under its common name in
many ‘non-taxonomic’ texts, including ‘conservation biology’ documents. Although
different scientific names have been used for this species in the recent years, it has been
clear to all involved that all those names designated the Aldabra tortoise and stabilisation
of its scientific name will be an easy task as soon as the Commission has made its decision.
In conclusion, we urge the Commission to refrain from using its plenary power to
suppress the holotype of Testudo gigantea Schweigger, 1812 or the name Testudo
dussumieri Gray, 1831, and to simply place both these names, as defined by their
name-bearing type specimens (respectively the holotype MNHN 9554 and the
lectotype RMNH 3231), as well as the generic names Aldabrachelys Loveridge &
Williams, 1957 and Dipsochelys Bour, 1982, as defined by their type species
(respectively Testudo gigantea Schweigger, 1812 and Testudo elephantina Duméril &
Bibron, 1835) on the Official Lists of Specific and Generic Names in Zoology.
Anonymous, 2001. Unesco world heritage convention periodic report. Part 11. Seychelles Island
Foundation. The Ministry of Environment, Seychelles: 1–40. [http://whc.unesco.org/
Beaver, K. & Gerlach, R. 1998. Aldabra management plan. Seychelles Island Foundation,
GEF – World Bank: 1–71. [http://whc.unesco.org/archive/periodicreporting/AFR/cycle01/
Boulenger, E.G. 1914. Reptiles and Batrachians. i–xiv, 1–278 pp. Dent & Sons, London.
Boulenger, G.A. 1889. Catalogue of the chelonians, rhynchocephalians, and crocodiles in the
British Museum (Natural History). i–x, 1–311, pl. 1–6 pp. Taylor & Francis, London.
Bour, R. 1982. Contribution à la connaissance des tortues terrestres des Seychelles: définition
du genre endémique et description d’une espèce nouvelle probablement originaire des îles
granitiques et au bord de l’extinction. Comptes rendus hebdomadaires des Séances de
l’Académie des Sciences de Paris, (3)295: 117–118, 121–122, 1 pl.
Bour, R. 1984. Taxonomy, history and geography of Seychelles land tortoises and fresh-water
turtles. Pp. 281–307 in Stoddart, D.R. (Ed.), Biogeography and Ecology of the Seychelles
Islands, Junk, The Hague.
Bour, R. 2006. Identity of Testudo gigantea Schweigger, 1812 and rediscovery of the type
specimen. Emys, 13(4): 12–23.
Dubois, A. 1997. Proposals concerning the conditions needed for a name being eligible for
conservation. Bulletin du Muséum national d’Histoire naturelle, (4)18(3–4): 317–320.
Dubois, A. 2005. Proposed Rules for the incorporation of nomina of higher-ranked zoological
taxa in the International Code of Zoological Nomenclature. 1. Some general questions,
concepts and terms of biological nameclature. Zoosystema, 27(2): 365–426.
Dubois, A. 2008. Identifying some major problems and their possible solutions. In Future trends
of taxonomy, EDIT Symposium, Carvoeiro (Portugal), 21–23 January 2008, Abstracts.
38–42 pp. Carvoeiro.
88 Bulletin of Zoological Nomenclature 67(1) March 2010
Dubois, A. & Raffaëlli, J. 2009. A new ergotaxonomy of the family Salamandridae Goldfuss,
1820 (Amphibia, Urodela). Alytes, 26(1–4): 1–85.
Duméril, A.M.C. & Bibron, G. 1834–54. Erpétologie générale ou Histoire naturelle complète des
Reptiles, 9 vols., 1 vol. pls. Roret, Paris.
Evans, L.T. & Quaranta, J.V. 1949. A study of dominance order in a herd of captive giant
tortoises (Testudinidae) resident at the Bronx zoo. P. 511 in American Society of
Zoologists, Forty-sixth annual meeting; New York City, December 28, 29, 30, The
anatomical Record, 105 (3).
Frazier, J. 2006. A neotype for the Aldabra tortoise, Testudo gigantea Schweigger, 1812.
Herpetological Review, 37(3): 368–373.
Fritsch, A. 1871. Zur Anatomie der Elephanten-Schildkröte (Testudo elephantina). Abhandlungen
der böhmischen Gesellschaft der Wissenschaften, ‘‘1870’’, (6), 4: 1–18.
Fritz, U. & Havaš, P. 2007. Checklist of Chelonians of the World. Vertebrate Zoology, 57(2):
Geiman, Q.M. & Wichterman, R. 1937. Intestinal Protozoa from Galapagos tortoises (with
descriptions of three new species. The Journal of Parasitology, 23(4): 331–347.
Georg, L.K., Williamson, W.L., Tilden, E.B. & Getty, R.E. 1962. Mycotic pulmonary disease
in captive giant tortoises, due to Beauvaria bassiana and Paecilomyces fumoso-roseus.
Sabouraudia, 2: 80–86.
Gerlach, J. 1997. Chelonia and people in Seychelles. Testudo, 4: 25–30.
Gerlach, J. & Canning, K.L. 1997. The Seychelles giant tortoise, its rediscovery and prospects
for conservation. Pp. 133–135 in Devaux, B. (Ed.), Proceedings of the International
Congress of Chelonian Conservation, Gonfaron, France, SOPTOM.
Günther, A. 1877. The gigantic land tortoises (living and extinct) in the collection of the British
Museum. i–v, 1–96, pls. 1–54. British Museum, London.
Peters, W. 1882. Naturwissenschaftliche Reise nach Mossambique. Zoologie. III. Amphibien.
i–xv, 1–191, pls. 1–26. Reimer, Berlin.
Pritchard, P.C.H. 1986. A reinterpretation of Testudo gigantea Schweigger, 1812. Journal of
Herpetology, 20(4): 522–534.
Quaranta, J.V. & Evans, L.T. 1949. Patterns of cooperative behavior in a herd of 14 giant
tortoises at the Bronx Zoo. P. 506 in American Society of Zoologists, Forty-sixth annual
meeting; New York City, December 28, 29, 30, The Anatomical Record, 105(3).
Rhodin, A.G.J., Parham, J.F., van Dijk, P.P. & Iverson, J.B. [Turtle Taxonomy Working
Group]. 2009. Turtles of the world: annotated checklist of taxonomy and synonymy, 2009
update, with conservation status summary. Chelonian Research Monographs, 5:
Schacht, P. 1903. Beiträge zur Kentniss der auf den Seychellen lebenden Elefanten-
Schildkröten. Pp. 103–129, pl. 15–21 in Chun, C., Wissenschaftliche Ergebnisse der
deutschen Tiefsee-Expedition auf dem Dampfer ‘‘Valdivia’’ 1898–1899, vol. III, Gustav
Schweigger, A.F. 1812. Prodromus monographiae Cheloniorum. Königsberger Archiv Naturwissenschaft
und Mathematik, 1: 271–368, 406–468.
Shea, G.M. 2001. Hyla lesueurii Bory de Saint-Vincent, 1828: An overlooked and problematic
frog species name. Journal of Herpetology, 35(2): 338–340.
Steers, J.A. 1968. Review of: Atoll Research Bulletin No. 118. The geographical Journal, 134(2):
Strauch, A. 1890. Bemerkungen über die Schildkrötensammlung im zoologischen Museum der
kaiserlichen Akademie der Wissenschaften zu St. Petersburg. Mémoires de l’Académie
impériale des Sciences de St.-Petersbourg, 38 (2): 1–127, 4 pls.
Urbain, A., Dechambre, E. & Piette, G. 1951. Septicémie due à Clostridium oedematiens, type
A, sur les tortues de la ménagerie des Reptiles du Muséum. Bulletin du Muséum national
d’Histoire naturelle, (2), 23(3): 247–248.
Wermuth, H. & Mertens, R. 1961. Schildkröten – Krokodile – Brückenechsen. i–xvi, 1–422 pp.
G. Fischer, Iena.