Comments on the type specimen of Dracaena guianensis Daudin,
1801 (Reptilia: Sauria: Teiidae), and rediscovery of the species in
French Guiana
J.-C. de Massary, M.S. Hoogmoed & M. Blanc
Massary, J.-C. de, M.S. Hoogmoed & M. Blanc. Comments on the type specimen of Dracaena guianensis Daudin, 1801 (Reptilia: Sauria: Teiidae), and rediscovery of the species in French Guiana.
Zool. Med. Leiden 74 (9), 15.ix.2000: 167-180, figs 1-8.— ISSN 0024-0672.
Jean-Christophe de Massary, Muséum national d’Histoire naturelle, Laboratoire de Zoologie (Reptiles
& Amphibiens), 25 rue Cuvier, F-75005 Paris, France. E-mail: massary@cimrs1.mnhn.fr
Dr. Marinus S. Hoogmoed, Nationaal Natuurhistorisch Museum, Postbus 9517, 2300 RA Leiden, The
Netherlands. E-mail: hoogmoed@nnm.nl
Michel Blanc, 24 les Jardins Sainte Agathe, 97355 Macouria, Guyane française.
Key words: Dracaena; holotype; rediscovery; type locality; range extension; French Guiana.
A discussion on the whereabouts of the holotype of Dracaena guianensis is given, and its rediscovery
in the National Museum of Natural History, Leiden, The Netherlands (RMNH) and the Muséum
national d’Histoire naturelle, Anatomie Comparée, Paris, France (MNHN-AC) is reported. The
RMNH possesses a stuffed specimen, of which the left forelimb has been amputated in life and of
which the skull is no longer present in the specimen. The Paris museum possesses a skull that fits the
Leiden specimen. Thus, the stuffed specimen and the skull together form the complete holotype, each
object can be considered a “schizotype” of the holotype. After one of us observed the species again in
French Guiana and documented this with a picture, the correction of the type locality to Amapá is
discussed and considered erroneous. Thus Cayenne again is considered the type locality.
Une discussion est menée à propos du holotype de Dracaena guianensis, et sa redécouverte au National Museum of Natural History, Leiden, The Netherlands (RMNH), et au Muséum national d’Histoire
naturelle, Anatomie Comparée, Paris, France (MNHN-AC) est indiquée. Le RMNH possède un spécimen empaillé, dépourvu de crâne, dont le membre antérieur gauche fut emputé lorsqu’il était encore
vivant. Le Muséum de Paris possède un crâne de cette même espèce. Après étude, il s’avère que le
crâne correspond tout à fait au spécimen empaillé ; ces deux pièces forment ensemble l’holotype, et
peuvent être considérées chacune comme un “ schizotype ” de l’holotype. En raison l’observation
récente de l’espèce en Guyane française, appuyé par un document photographique, la correction de la
localité type à l’Amapá est discutée, et considérée comme éronnée. Cayenne est à nouveau considérée
comme la localité type.
Uma discussão sobre o holotipo de Dracaena guianensis é apresentada, registrando-se sua redescoberta
no Nationaal Natuurhistorisch Museum, Leiden, Países Baixos (RMNH), e no Muséum national
d’Histoire naturelle, Anatomie Comparée, Paris, França (MNHN-AC). O RMNH possui um espécime
empalhado, destituído de crânio, no qual o membro anterior esquerdo foi amputado quando o animal
ainda vivia. O MNHN-AC possui um crânio dessa mesma espécie. Nossos estudos demonstram que o
crânio corresponde ao espécime empalhado, os quais em conjunto formam o holótipo de Dracaena
guianensis. Crânio e espécime constituem, portanto, cada um, um “schizótipo”. Tendo em vista a
observação recente da espécie na Guiana Francesa por um de nós, documentada por foto, rejeita-se a
correção da localidade-tipo como sendo o Amapá. Cayenne volta a ser a localidade-tipo.
Introduction
Two species are presently recognised in the lizard genus Dracaena. The most widespread species, Dracaena guianensis Daudin, 1801, occurs throughout Amazonia, essentially following the Amazon river and some of its principal (southern) tributaries
(Avila-Pires, 1995: 546). The second species, Dracaena paraguayensis Amaral, 1950, is
168
Massary et al. Comments on Dracaena guianensis. Zool. Med. Leiden 74 (2000)
restricted to Mato Grosso do Sul, Brazil and Paraguay (Vanzolini & Valencia, 1965).
D. guianensis is a large and stout teiid lizard reaching a maximum snout-vent
length of 360 mm (Avila-Pires, 1995: 541). It lives in swampy areas and spends most
of its time on low trees, though feeding only occurs in water (Vanzolini & Valencia,
1965); it feeds mostly on shells which it breaks with its strong molariform posterior
teeth. Because of its ecological habits and its skin resembling that of crocodilians
(caimans), it is commonly named “caiman-lizard” in English and “lézard-caïman” in
French; La Cepède (1788) called it “La dragonne”: he was also the first to give a good
description and a picture of this species. Unfortunately, he used the name Lacerta
dracæna [sic], previously used by Linnæus (1758) for a varanid species (see Opinions
International Commission on Zoological Nomenclature, 1959). Thus, La Cepède’s
description was without available name. In fact, Daudin (1801) was the first to give a
good description and an acceptable binomen, Dracaena guianensis, still used today.
The aim of this note is to update the situation concerning the type specimen and
its probable locality in the light of recently acquired data.
Discussion on the type specimen
When describing “La dragonne”, La Cepède (1788) gave details for only one specimen. It is clear he only had one specimen and not more from the fact that in the
description he always used the singular, and especially the following part of a sentence leaves no doubt about the material he saw (La Cepède, 1788: 245): “ …il y a, au
Cabinet du Roi, un individu de cette espèce, qui a été envoyé de Cayenne par M. de la
Borde, & d’après lequel nous avons fait la description que l’on vient de lire (c); …..”.
The footnote (c) refers to a table of measurements of the specimen, which was again
repeated by Daudin (1801). Later, Daudin (1801: 425) wrote about D. guianensis: “Je
regarde donc comme vrai type de dragone [sic] un saurien qui a été envoyé, il y a
quelques années, de Cayenne au museum [sic] d’histoire naturelle de Paris par
Delaborde [sic], et que le savant professeur Lacépède [sic] a décrit et figuré dans son
ouvrage sur les quadrupèdes ovipares”. From this sentence it is clear that the type
specimen for the binomen Dracaena guianensis Daudin, 1801 is the same specimen as
that described by La Cepède in 1788 as “La dragonne”. In addition, Daudin (1801:
429) gave a table with exactly the same meristic data as provided by La Cepède (1788:
245). The legend of this table reads as follows: “Dimensions de la dragone [sic] qui est
conservée dans la collection du museum [sic] d’histoire naturelle de Paris, Par
Lacépède [sic]”. Thus, Daudin clearly indicated that he saw the only specimen of Dracaena guianensis available at that time, and which already had been described by La
Cepède (1788).
MNHN 8735 has been considered the type specimen of Dracaena guianensis (e.g.
Hoogmoed & Lescure, 1975: 166; Brygoo, 1989: 19; Avila-Pires, 1995: 540). In fact, this
assumption is wrong. The size data given by La Cepède (1788: 245), and repeated by
Daudin (1801: 429), do not agree with those of MNHN 8735 (Table 1). For instance,
the La Cepède’s specimen had a total length of 790 mm1, against 925 mm for MNHN
1
All the measurements provided by La Cepède (1788) and Daudin (1801) are in ancient French units
as follows: 1 pied = 325 mm, 1 pouce = 27.07 mm, 1 ligne = 2.25 mm. The measures cited in the present paper were all converted to mm.
Characters
La Cepède 1788
Head length
Neck length
Trunk length
Tail length, total
Total length
Mouth outline
Distance between eyes
Maximum body circumference
Tail base circumference
One limb absent
Hind limb
Fore limb
Skull/head width, widest point at temple
Skull/head width, posterior part skull
Skull/head length to posterior edge tympanum
Skull/head length to anterior edge tympanum
Snout-vent length
Tail regenerated part
Skull length (greatest length), left
Skull length to end auditory bulb, left
Skull/head width below eyes
Skull/head width base skull betw auditory bulbs
Skull height
Interorbital distance
Length left mandible
Length right mandible
?
?
?
447
794
117
27
203
153
yes
148
103
-
Daudin 1801
RMNH 28888
Dum & Bibr 1839
MNHN 8735 in 1999
skull MNHN-AC
2000.242
?
?
?
447
794
117
27
203
153
?
148
103
-
see below
44
195
440
755
135
26.8
220
145
yes
bent
bent
51.3*
46.4*
76*
69.1*
315
180
38.3*
47.5*
8.5*
-
90
60
220
570
790 (instead of 940)
?
?
?
?
?
170
130
-
85
60
210
560
925
175
29
245
160
no
165
128
75.7*
104*
112*
365
60.3*
65.2*
56.1*
-
34
41.4
80
69.2
83
81.6
37.9
41.4
26.2
8.5
83.0
83.8
Massary et al. Comments on Dracaena guianensis. Zool. Med. Leiden 74 (2000)
Table 1.— Table comparing characters given for the specimens of Dracaena guianensis studied by La Cepède (1788), Daudin (1801) and Duméril & Bibron
(1839), for the stuffed specimen in Leiden (RMNH 28888), for the Paris skull and mandibles (MNNH-AC 2000.242) and for MNHN 8735, erroneously considered as Daudin’s holotype for Dracaena guianensis. Note that the total length given by Duméril & Bibron (790 mm) for this latter specimen does not correspond to the sum (940 mm) of all other parameters. (Question marks = lacking data; - = measurements could not be taken; * indicates external measurements
of the head, not the skull; all measures in mm).
169
170
Massary et al. Comments on Dracaena guianensis. Zool. Med. Leiden 74 (2000)
8735. Moreover, La Cepède (1788: 19) in a footnote to the general part of the book, in
a discussion on regeneration, far removed from the description of “La dragonne”,
revealed an additional, notable character: “L’on conserve au Cabinet du Roi, un
grand lézard, de l’espèce appelée dragonne, auquel il manque une patte [= one limb
is missing]; il paraît qu’il l’avait perdue par quelqu’accident, lorsqu’il était déjà assez
gros; car la cicatrice qui s’est formée est considerable. C’est M. de la Borde, Médecin
du roi à Cayenne, & correspondant du Cabinet du Roi, qui l’a envoyé”. Unlike the
specimen studied by La Cepède and sent by La Borde from Cayenne (French Guiana),
MNHN 8735 has four limbs, although some fingers and toes now are missing. We
therefore come to the conclusion that MNHN 8735 certainly can not be the specimen
described by La Cepède in 1788.
In the dry collections of the Nationaal Natuurhistorisch Museum (formerly
Rijksmuseum van Natuurlijke Historie) (acronym to be used RMNH) one of us
(MSH) had noted in the late nineteensixties a stuffed specimen of Dracaena guianensis
(RMNH 28888) which was rather peculiar because it lacked the left forelimb, had all
the correct label data corresponding with those given by La Cepède (1788), but never
was considered the specimen described by La Cepède (1788), because in the description La Cepède never mentioned the very obvious fact that one limb was missing.
Now that the senior author of this paper (JCM) discovered the remark in the footnote
on p. 19 (La Cepède, 1788), mentioned before, about the specimen missing a leg,
things suddenly became clear and we now can safely conclude that the RMNH specimen is the one that was used by La Cepède (1788) for his desciption and later on also
by Daudin (1801). The specimen is a stuffed specimen, mounted on some wooden
planks. It is accompanied by the following data:
A label which has been nailed to the side of the main plank, reading in bold lettering: Crocodilurus lacertinus (which has been striked out and replaced by “Dracaena
guianensis Daud” in smaller and more delicate letters), Mus. Paris , Cayenne (fig. 1).
There are more data are on the bottom of the planks as follows:
A cardboard label nailed to the central, longitudinal plank, that reads (words
written in ink):
Crocodilurus dracaena
La dragonne
Mus. Paris Cayenne
The (obviously handmade) label has a red edge, composed of an outer red line,
separated by some white from six very closely arranged red lines forming a band,
and an inner red line, again separated by some white from the central red band.
These red ink lines have been drawn by hand (fig.4).
The text on the label is also written on the white-painted part of the underside of
the same central plank in a different handwriting and in pencil (fig. 5).
Partly hidden by the label there was some more text on two areas that were not
painted white (as most of the central plank is). Part of these texts (in ink) is partly
covered by smears of white paint (fig. 5).
In the left unpainted area one can read (all in ink):
Dragone de la guyane
D. guyanensis
envoye de Cayenne
Massary et al. Comments on Dracaena guianensis. Zool. Med. Leiden 74 (2000)
171
Fig. 1. RMNH 28888, Dracaena guianensis Daudin, 1801. From top to bottom left side view, dorsal view and right side view of stuffed holotype.
172
Massary et al. Comments on Dracaena guianensis. Zool. Med. Leiden 74 (2000)
par M° de la Borde
The first letter of Cayenne looks like a capital G was first written, which subsequently was changed into a capital C (fig. 5).
In the right unpainted area one can read (all in ink, except where noted differently):
La grande Dragonne Cuv.
La Dragon Lacep. Quad. Ovip. pl.IX
Tejus [in pencil]
Monitor crocodilus Merr.
Mus. a Paris
1835
To the left of the text there is a large sign which looks like a capital letter S leaning
backwards, but turned 90° looks like a capital letter L, written in pencil. If we would
interpret this sign as a capital S indeed, it looks completely different from the way
Schlegel, who worked in the RMNH between 1825 and 1884 and in 1835 was responsible for the herpetological collections, wrote a capital S. So it probaly is not his handwriting (fig. 5).
Under the plank used to support the hindlimbs, under the left hindlimb the number 84 is written in pencil. Possibly meant to be a register number, but never effectuated as such in the present RMNH collections and registernumber system (fig. 5).
In the archives of the RMNH one of us (MSH) found a list, signed by H. Schlegel
and dated June 1835, detailing the material that was received in Leiden as exchange
from the Paris museum. Under number 68 is listed “La grande Dragonne de Lacep.
(ind[ividu] emp[aillé]).
All these data combined make it very likely, rather undeniable, that the stuffed
specimen in Leiden (RMNH 28888) is the original specimen on which La Cepède
(1788) based his description of “La Dragonne”.
The measurements taken from the available material and compared with data
from the literature (table 1) show that RMNH 28888 in most measurements indeed is
close to, or identical with, the measurements of the specimen described (and measured) by La Cepède (1788) and Daudin (1801). Also, some head measurements of
Fig. 2. RMNH 28888, Dracaena guianensis Daudin, 1801, holotype, detail of head and left side of the
body, showing the scar at insertion of left forelimb.
Massary et al. Comments on Dracaena guianensis. Zool. Med. Leiden 74 (2000)
173
Fig. 3. RMNH 28888, Dracaena guianensis Daudin, 1801, holotype, detail of scar at insertion of left forelimb.
RMNH 28888 closely agree with those of the skull (MNHN-AC 2000.242) (see below).
RMNH 28888 is a stuffed specimen (fig. 1) which lacks the skull and the axial skeleton, only the bones in the limbs are still present (checked by X-raying the specimen).
The body and limbs are supported by thick ironwire, with which the specimen also is
attached to the planks on which it is mounted. It is in relatively good condition, but
some damage is evident: the right forelimb is complete, all fingers are present; the left
foot lacks the 2nd and most of the fifth toe; the right foot lacks toes 1, 2, 3 and 5, and
most of 4; the tail has been broken (and repaired again) at cm 19.5 and there the filling
material is visible over a distance of 4 to 5 mm; the shape of the head is rather asymmetrical and irregular, due to the fact that the skull has been removed; the mouth on
both sides is slightly opened and shows filling material; the eyes have been replaced
by round beads, the one on the left is quite visible, the one on the right is covered by
the lower eyelid. As stated before, the left forelimb is missing, due to some accident
that happened to the animal while still alive, as the scar where the limb has been severed is neatly healed with skin, showing more or less radially arranged small scales
(granules) of slightly variable size. The scar has a diameter of 16.3 mm and agrees
with the general description of it given by La Cepède (1788) (figs 2, 3). The tail is
regenerated from cm 26.5 , a character also recognisable in the drawings published by
La Cepède (1788, 1799) and Daudin (1801). From the fact that this specimen was
exchanged with the Leiden museum we might conclude that at the time of exchange
the Paris museum already had a second specimen of the species, viz. MNHN 8735.
The fact that the drawing of La dragonne in La Cepède (1788) seems to show an animal facing left, with a left forelimb, does not mean much, as the picture probably was
174
Massary et al. Comments on Dracaena guianensis. Zool. Med. Leiden 74 (2000)
Fig. 4. Red-edged label attached to bottom of central plank on which the stuffed holotype of Dracaena
guianensis Daudin, 1801 (RMNH 28888) is mounted.
printed in mirror-image as happened often. This is made more likely by the fact that
in the 1799 edition of La Cepède, published in Dordrecht, the same plate is used
again, but this time it is printed the other way, the animal facing right and apparently
showing a right forelimb. Also Daudin (1801) gives a picture in which the animal is
facing right, and showing a right forelimb. In all pictures two hindlimbs are recognisable, but only one forelimb, which might possibly reflect the fact that the left forelimb
is absent.
Now that it has become clear that RMNH 28888 lacks a skull, the fact that a skull
(MNHN-AC 2000.242) prepared and drawn by George Cuvier (1824), was found in
the Laboratoire d’Anatomie comparée du Muséum national d’Histoire naturelle,
becomes extra interesting. The skull (figs 6, 7) was sent from Paris to Leiden for a
direct comparison with the stuffed specimen. Already at first view it was clear that
the skull matched the stuffed specimen very well and this was corroborated by the
measurements taken (table 1). The skull is slightly damaged in the occipital area: the
posterior part of the parietal on the left hand side is missing (cut off along an oblique
line, from anterolateral to posteromedial); on the left hand side the posterior part of
the postocular and the squamosal are missing; on the right hand side the postocular
and part of the parietal (small chip in the middle) are missing. The premaxillary teeth
are small and pointed, the anterior five or six maxillary teeth are also small and pointed, the posterior five maxillary teeth are enlarged and rounded, with the last one very
much smaller than the preceding four. The first four mandibular teeth are small and
pointed, teeth 5-7 are intermediate and 8-12 are flattened, with the last one distinctly
smaller than the preceeding four.
Thus, La Cepède’s original specimen still exists and now is divided over two
museums: the stuffed body with the bones of the limbs still in it in Leiden (RMNH
28888) and the (slightly damaged) skull and the two mandibles in Paris (MNHN-AC
2000.242). Together these parts form the holotype of Daudin’s Dracaena guianensis and
could be considered “schizotypes”, a rather unusual occurrence in herpetological
specimens.
Massary et al. Comments on Dracaena guianensis. Zool. Med. Leiden 74 (2000)
175
Fig. 5. RMNH 28888, Dracaena guianensis Daudin, 1801, holotype, view of tekst on bottom of planks on which stuffed specimen is mounted, after removal of
the red-edged label shown in fig. 4.
176
Massary et al. Comments on Dracaena guianensis. Zool. Med. Leiden 74 (2000)
Fig. 6. Ventral and right lateral view of skull and interior view of mandibles of the holotype of Dracaena guianensis Daudin, 1801 (MNHN-AC 2000.242) kept at the Laboratoire d’Anatomie comparée du
Muséum nationale d’histoire naturelle de Paris.
Massary et al. Comments on Dracaena guianensis. Zool. Med. Leiden 74 (2000)
177
Fig. 7. Dorsal view of skull and mandibles of the holotype of Dracaena guianensis Daudin, 1801
(MNHN-AC 2000.242) kept at the laboratoire d’Anatomie comparée du Muséum nationale d’histoire
naturelle de Paris in a box with skulls prepared by Cuvier (photo J.-C. de Massary).
On the origin of MNHN 8735
Duméril & Bibron (1839: 59) gave a total length of 790 mm for the specimen they
described ; but this is inconsistent with the other size data they provided. Indeed, as
noted first by Brygoo (1989: 39), the addition of the different measurements given by
Duméril and Bibron, head plus neck plus trunk plus tail, yields 940 mm and not 790
mm. These latter authors probably omitted to take into account the head and neck
length so that they would only get 790 mm. Except for the total length, all the measures we took on MNHN 8735 correspond well with the size data presented by
Duméril & Bibron (Table 1) so that we do not doubt that this specimen was the one
described by these authors.
Duméril & Bibron (1839: 60) also indicated “Nous avons peu de choses à dire sur le
mode de coloration de cette espèce, dont nous n’avons jamais vu que des individus
desséchés”; from this phrase, we conclude that they saw more than one specimen, but
we do not know exactly how many ; further down the text, they added “le Muséum
d’histoire naturelle l’a anciennement reçu de la Guyane par les soins de M. de Laborde
[sic]; mais depuis, elle ne s’est jamais trouvée dans aucune des collections qui nous ont
été adressées de ce pays.” which means that, whatever the number of specimens they
had, they apparently were of the opinion these were all sent by La Borde from
Cayenne. Apparently they did not see any specimens in other shipments from French
Guiana. Therefore, we could assume that MNHN 8735 also came from French Guiana
178
Massary et al. Comments on Dracaena guianensis. Zool. Med. Leiden 74 (2000)
and was (possibly) also sent by La
Borde, just like the type specimen. We
only could ascertain that Raymond de
La Borde (or Raymond de Laborde),
born in 1725, arrived in French Guiana
on March 18, 1769 on board of the ship
“L’Amphitrion” that arrived from Bordeaux, France and stayed there at least
till 1776. There apparently is some
information indicating he still was in
French Guiana in 1782. He died in 1786
(possibly in French Guiana). No other
information on him is available to us at
Fig. 8. Live Dracaena guianensis (not collected), 2
the moment. From the scant informajune 1996, at 4 meters height in a bush, Kaw river,
tion above it is however, certain that
French Guiana. (photo M. Blanc).
La Borde could not have collected any
material in French Guiana after 1801
(see below).
The historic data available, enable us to make a rough estimate of the period in
which MNHN 8735 could have arrived in the Paris museum. We know that both La
Cepède and Daudin saw only one (and the same) specimen; we thus can conclude
that MNHN 8735 has arrived at the Paris Museum after 1801, the date of publication
of the second volume of the Histoire naturelle, générale et particulière, des reptiles
by Daudin. From the fact (see above) that the specimen described by La Cepède was
exchanged with the Rijksmuseum van Natuurlijke Historie in June 1835, we may
safely assume that at that time the Paris museum had more than one specimen and
that the “least valuable” one (damaged, because it was missing one forelimb) was
used for exchange and that MNHN 8735 was present at that time and thus had
arrived somewhere between 1801 and June 1835, and could not have been collected
by La Borde (see above).
In 1851, Duméril & Duméril reported only one specimen, the actual MNHN 8735,
in the museum collections; this specimen remained the only complete Dracaena
known in the Paris Museum collections until 1973.
On the type locality
Contrary to what Hoogmoed & Lescure (1975: 166) thought, Duméril & Bibron
(1839: 60) did not doubt the origin of the La Borde’s specimen; Duméril & Bibron just
noted that this species was only known from Cayenne thanks to La Borde, that is all.
The last part of their sentence (already cited above) established that the species was
absent in other shipments (not sent by La Borde) from French Guiana. Because the
ancient southern limit of French Guiana was the Rio Araguari and because there was
no evidence of the occurrence of Dracaena guianensis within the present borders of
French Guiana, Hoogmoed & Lescure (1975) logically concluded that the type locality
“Cayenne“ very probably was wrong; consequently, they corrected it to Amapá,
north of the Rio Araguari, Brazil.
Massary et al. Comments on Dracaena guianensis. Zool. Med. Leiden 74 (2000)
179
However, one of us (MB) recently took a picture of Dracaena guianensis in the wild
at the Kaw river, north-eastern French Guiana, where the species is not very rare.
Though the quality of the picture is not very good, the lizard in this picture (fig. 8)
clearly can be recognised as a caiman-lizard; indeed, the dorsal heterogenous lepidosis and the large dispersed tubercles recalling caiman skin can easily be seen. Also the
typical head shape, laterally compressed and flat above and the reddish colour of the
head are recognisable.
Because of the ancient limits of the French Guianan territory, we cannot be
absolutely certain that the specimen sent by La Borde to La Cepède was caught inside
the actual limits of French Guiana; yet, in the light of the recent observation we cannot reject this possibility any longer. Therefore, the type locality correction made by
Hoogmoed & Lescure (1975) can be repealed and we have to accept again Cayenne (=
French Guiana) as the type locality of Dracaena guianensis Daudin, 1801. The recent
picture also means that the distribution of the species is more extensive than thought
before and that it is now possible to say that it occurs throughout Amazonia, essentially following the Amazon river and some of its principal (southern) tributaries, but
in the northeastern part of its distribution extends its range north throughout
(coastal) Amapá and reaches north-eastern French Guiana. A similar distribution has
become known the past few years for other reptiles and amphibians as well.
Acknowledgements
We want to thank Dr T.C.S. Avila Pires for providing the Portuguese resumen.
Figures 1–6 were made by Mr A. ‘t Hooft of the RMNH. We would like to thank Dr J.P. Gasc (MNHN-AC = Muséum national d’Histoire naturelle, Anatomie Comparée,
Paris) for loaning the MNHN skull and mandibles to MSH, thus enabling direct comparison with the RMNH stuffed specimen. The stuffed specimen RMNH 28888 was
X-rayed by Mr P. Snelderwaard of the Zoological Laboratory (v. d. Klaauw Laboratory) of the Leiden University. JCM wants to thank Dr J. Lescure and Dr A. Dubois for
stimulating discussions on the subject. He also wants to thank Dr J.-P. Gasc and Dr I.
Ineich for reviewing an earlier draft of this paper. Mrs Monique Pouliquen of the central library of the MNHN provided biographic information on La Borde. The taxonomic work for this paper took place complementary to an ecological study on
lizards led by JCM in French Guiana, and supported by Electricité de France (EDF)
(convention EDF/MNHN GP 7531). MNHN is the acronym for the general collection
of the Muséum nationale d’Histoire naturelle, Paris, France, and RMNH for National
Museum of Natural History, Leiden, The Netherlands.
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Received: 17.iii.2000
Accepted: 4.iv.2000
Edited: R. de Jong