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New light on an old tortoise - Gilbert White's Selborne tortoise re-discovered.
A C Highfield & J. Martin
TESTUDO WHITEI BENNETT 1836 SUMMARY
N. African members of the genus Testudo have in recent years all
been regarded as belonging to the same species, Testudo graeca L.
1758. This paper describes a form which occurs in Algeria,
geographically immediately adjacent to the race recognised as T. graeca
L. 1758, but which demonstrates sufficient divergence of
characters to require separate classification as an autonomous
species. Indeed, it has so been described previously (Bennett, 1836),
but in subsequent revisions of the genus (Wermuth and Mertens 1961,
Loveridge and Williams, 1957) has been erroneously relegated as a
synonym of T. graeca L., Testudo ibera (Syn. T. graeca L., Boulenger
1889) or of Testudo marginata SCHOEPFF 1792 (Gray 1870). The object of
this present paper is to demonstrate that its earlier status as a
separate species was justified and should therefore be restored.
KEYWORDS: Testudo graeca - Gilbert White - Algeria - Taxonomy - Size -
Age - Nomenclature
INTRODUCTION
The Testudo graeca of Linnaeus (1758) is based upon the holotype of
Edwards (1748) (Terra typica = Santa Cruz, Oran, Algeria). An
excellent illustration of this carapace is provided by Edwards (1748)
P.204. Synonyms are proliferate, good summaries of which are
provided by Loveridge & Williams (1957), Siebenrock (1910), Strauch
(1862) and Boulenger (1891). It should be noted that many
illustrations purporting to show T. graeca L. 1758 actually depict
other races;- two such examples are Sowerby & Lear's plate (1872)
which although labeled Testudo graeca is unmistakably that of a
Testudo ibera PALLAS 1814 and Lortet's (1888) fine plate of a
Testudo hermanni GMELIN 1789 which is mis-titled Testudo graeca L.
Most published descriptions of land tortoises in Algeria are
unfortunately rather unsatisfactory. Compound descriptions are
particularly commonplace, with regular confusion between the race T.
graeca L. 1758 , T. ibera PALLAS 1814, T. marginata SCHOEPFF 1792
and T. hermanni GMELIN 1789 which was habitually referred to as T.
graeca L. by many authors of the period. Few 19th century authors
were sufficiently familiar with the true diagnostic characters of
these various races and both synonyms and composite descriptions are
therefore endemic in their writings e.g, see Gervais (1836, 1848 and
1857), Strauch (1862), Doumergue (1904) Tristram (1860) and Lataste
(1880 and 1881).
Geographically erroneous reports of T. marginata in Algeria may
however have significance in the light of our present findings. It
seems quite possible that the basis of these reported sightings could
have been the large Testudo spp. described and illustrated here.
These must have been encountered in the field, but seem to have been
explained as simply elderly specimens of T. graeca L. which had
attained an unusual size e.g, writing of Testudo graeca Boulenger
(1891) comments;
"Old specimens have been taken for the allied T. marginata Shoepff,
s. (syn) campanulata STRAUCH (by Gervais and Lallemant), the habit
of which appears to be restricted to Greece".
This question of confusion regarding the distribution of T. marginata
is also discussed by Lortet (1888) who states that he received many
reports of marginata from Algeria, but felt that they were not
accurate, being based upon specimens of T. ibera PALLAS (syn. T.
mauritanica Dumeril et Bibron) which may "possess similar characters"
(p.9). It should be noted that M. Lortet's usage of T. ibera PALLAS is
itself synonymous with Testudo graeca L. as is Dumeril and Bibron's
Testudo mauritanica (Dumeril and Bibron, 1835).
Gilbert White's "Timothy" Tortoise of Selborne
Readers of Whites classic work The Natural History of Selborne (1789)
cannot fail to have been entertained by his vivid descriptions
of the activities of the "old Sussex tortoise" which passed to him
following the death of his aunt in 1780. It had been purchased by his
uncle Mr. Snooke from a sailor in Chichester for 2/6d sometime during
1740. White knew the tortoise prior to obtaining possession of it from
his many visits to his relatives who resided at Delves House,
Ringmer, where the animal lived (Wright, 1917). As White explains
in one of his letters to Daines Barrington dated 8th October 1770;
" A land tortoise, which has been kept for thirty years in a little
walled court belonging to the house where I am now visiting, retires
under the ground about the middle of November, and comes forth again
about the middle of April. When it first appears in the spring it
discovers very little inclination towards food; but in the height of
summer grows voracious: and then as the summer declines its appetite
declines; so that for the last six weeks in autumn it hardly eats at
all. Milky plants such as lettuces, dandelions, sowthistles are its
favorite dish. In a neighboring village one was kept till by tradition
it was supposed to be an hundred years old. An instance of vast
longevity in such a poor reptile!".
Another letter dated 21st April 1780 continues;
"The old Sussex tortoise, that I have mentioned to you so often, is
become my property. I dug it out of its winter dormitory in March last,
when it was enough awakened to express its resentments by hissing; and,
packing it in a box with earth, carried it eighty miles in
post-chaises.... as it will now be under my eye, I shall now have an
opportunity of enlarging my observations on its mode of life, and
propensities".
This White certainly did, recording assiduously the behaviour of
"Timothy" until a few days before he died in 1793. A collection of
Whites notes concerning the tortoise has since been published
separately (Warner, 1946). An excellent summary of Gilbert White's
involvement with 'Timothy' tortoise and a brief history of Selborne can
also be found 'Selborne and the Tortoise Connection' (Chatfield, 1986).
Of precise data concerning White's companion we have some useful
records. We note that over the 18 years White had charge of it, its
weight fluctuated between 2,835g to 3,260g. Its carapace (which is now
preserved in the British Museum collection) measures 250mm. White
himself did not realise that it was a female, nor did he know its
origin, for he speculates that it may have come from Virginia, U.S.A.
He did, however make a reasonable guess at what it might have been;
"Timothy seems to be the Testudo graeca of Linnaeus" (Journal of
Gilbert White, July 1st 1780).
White's confusion in respect of Virginia may well have some foundation
in the fact that his aunt, Mrs. Snooke, also kept Box Tortoises
(Terrapene spp.) which originate in the U.S.A.
The Torquay Tortoises
Flower (1945) reported in some detail on a pair of female
tortoises of Algerian descent which were subsequently maintained in
captivity in England for 39 years. Flowers excellent report, based
upon records kept by their owner, a Mr. Moysey of Torquay, chronicles
the growth of the pair from 1906-7 when they weighed and measured
0.012kg/49mm and 0.008kg/47mm (measured over the curve of the
carapace) respectively to 1944 when their measurements were
3.969kg/365mm and 1.814/279mm. A further set of data on the same
tortoises was presented by Loveridge and Williams (1957) where it is
recorded that as of September 1st 1955 the tortoises weighed
2.629kg/298mm and 4.252kg/365mm and by F. E. Moysey (1962). The largest
of the pair died in 1969 at the age of 63 years having attained a
maximum weight of 4.366kg (Lambert, 1982).
The parents of these tortoises were collected in the Foret de Bainen a
few kilometers west of Algiers. Eggs were laid during May 1905,
and from the 14 hatchlings "Daimler" and "Panhard" were
eventually selected and bought to England. The illustration
accompanying Flowers report clearly identifies them as belonging
to the same race as the specimens in plate 1 and not as T.
graeca L. 1758 as Flowers believed. Flowers was however correct
in his conclusion that "very large" tortoises "do not necessarily
come from Syria, Asia Minor, or the Balkan Peninsular, but may be
from the typical Linnean locality of north-west Africa."
The claim that this form of Algerian Testudo, as depicted by
Flower (1945) are comprised of nothing more than extremely old or
unusually large T. graeca L. is not supported by the evidence.
This form, henceforth referred to as Testudo whitei BENNETT 1836,
consistently exhibits entirely different carapace markings from
T. graeca L.
Both T. graeca L. and T. whitei attain full adult
dimensions at approximately the same age, typically between 25-35
years of age. Testudo whitei however consistently and invariably
attains a much greater size than that ever recorded for specimens
conforming to the holotype of T. graeca L. After approximately 40 years
of age, growth in both forms slows very considerably eventually
becoming almost imperceptible.
The plastral markings of the two races are consistently different.
The general carapace morphology of T. whitei is quite different from
that of T. graeca L. with each race demonstrating an entirely
different height-length ratio.
COMPARISON OF CHARACTERS
Carapace markings: T. graeca L. 1758
Groundcolour amber-yellow to greenish-yellow, dark brown to black
central dot on vertebral and costal scutes, vertebral and costal
scutes feature distinctive brown-black lateral "ring" marking, the
central marginal scutes feature a distinctive brown-black triangular
"saw-tooth" pattern. The plastron typically features 3 pairs of large
brown-black dots (plate 2).
Carapace markings: Testudo whitei BENNETT 1836
Groundcolour light amber-yellow, less distinctive central brown-
black markings on vertebrals and costals, sometimes entirely absent.
No lateral "ring" markings, very distinctive medullary "ray" pattern
radiating from areolae on vertebral and costal scutes usually
present. Some specimens exhibit fewer markings on an almost
unbroken even yellow-amber ground colour. The characteristic dark
brown-black triangular marginal markings of T. graeca L. are
invariably either absent entirely or ill defined compared to T.
graeca L. The plastron features a larger and more diffuse expanse of
black markings than T. graeca L. (plate 3).
Carapace morphology and size: T. graeca L. 1758
The following measurements were taken from six randomly selected adult
females conforming to the holotype of T. graeca L. as illustrated in
plates 4 & 5. The age range of the individual animals was estimated at
between 35 years to 85 years. Some specimens had been in captivity in
England for over 60 years.
Length |
Carapace height |
Maximum width |
Weight |
192mm |
104mm |
138mm |
1,500g |
190mm |
99mm |
133mm |
1,250g |
187mm |
105mm |
135mm |
1,300g |
176mm |
94mm |
125mm |
1,000g |
165mm |
95mm |
119mm |
800g |
153mm |
84mm |
109mm |
790g |
The specimen at 192mm is the largest female example of this race
measured to date.
Carapace morphology and size : Testudo whitei
The following series of measurements were obtained from eight adult
Algerian females conforming to the type of T. whitei and examined
recently by the authors (plates 1 and 6). The age range of the animals
was estimated at between 35 years to 85 years. The specimen measuring
270mm was known to be less than 40 years of age with reasonable
certainty, having been in captivity for over 30 years and from a time
when it measured less than 70mm long.
Length |
Carapace height |
Maximum width |
Weight |
280mm |
134mm |
182mm |
3,750g |
280mm |
125mm |
190mm |
3,575g |
272mm |
127mm |
130mm |
3,400g |
270mm |
130mm |
180mm |
2,875g |
268mm |
135mm |
180mm |
3,300g |
250mm |
125mm |
190mm |
3,000g |
245mm |
120mm |
165mm |
2,600g |
240mm |
118mm |
165mm |
2,200g |
Comparative morphology female T. graeca L. vs. T. whitei BENNETT
Apart from the obvious difference in maximum dimensions and body mass,
if a length-height index is prepared by dividing the straight line
carapace length by the maximum carapace height the difference in body
shape is also evident, the female T. graeca L. having a relatively
higher domed carapace than those of T. whitei BENNETT which are
broader and flatter. The indices obtained from the eight examples of T.
whitei described above are 2.09, 2.24, 2.14, 2.08, 1.99, 2.00,
2.03 and 2.04. If the same formulae is applied to the six female T.
graeca L. the results are; 1.85, 1.92, 1.78, 1.74, 1.87 and 1.82 - an
altogether lower average index for the race which is consistent across
all examples examined to date.
Whilst these measurements were being made, it was further observed that
female specimens of T. graeca L. generally exhibited a much wider
posterior marginal flare than the measurement of the width of the body
at the median line. This character is not typical of Testudo whitei,
where the median carapace width of females often is equal to or exceeds
the maximum lateral width attained by the posterior marginals. In only
one of the eight specimens of T. whitei detailed above was the
posterior marginal maximum width greater than that of the body at the
transverse median line (the specimen measuring 272mm) and in this case
it was noted that the tortoise was unusually elongate and narrow in
form. The median width was 125mm, yet the maximum marginal width was
recorded at 164mm. This latter measurement is not unusual, but the
narrowness of the waist in this specimen in our experience is most
unusual. In all other respects the specimen was absolutely normal.
Sexual dimorphism male Testudo graeca L. & Testudo whitei BENNETT
The males of both species attain considerably smaller dimensions than
the females. The dimensions of the two largest specimens of each race
we have seen to date are as follows;
Testudo whitei BENNETT 1836
Length |
Carapace height |
Maximum width |
Weight |
240mm |
111mm |
154mm |
2,350g |
Length-Height ratio = 2.18
Testudo graeca L. 1758
Length |
Carapace height |
Maximum width |
Weight |
188mm |
102mm |
118mm |
1,150g |
Length-Height ratio = 1.84
Most specimens are much smaller than this, the two above tortoises
being unusually large and somewhat exceptional examples of their
respective species. The different H-L ratios express the lower domed
and much more elongate carapace of males belonging to the race T.
whitei compared to males of T. graeca L. The following tables provide
typical comparative length-weight data for males of the two species;
Testudo graeca L. |
Testudo whitei BENNETT |
Length |
Weight |
Length |
Weight |
155mm |
750g |
223mm |
2,200g |
145mm |
675g |
215mm |
1,750g |
160mm |
800g |
220mm |
2,000g |
It will be noted that males of T. whitei typically weigh over 1kg more
than males of T. graeca L. The figures of weights and lengths obtained
from males should be compared to the data taken from females. In both
races, males are typically considerably smaller than females with a
lower body mass and H-L ratio representing the reduced 'dome' or
carapace curvature.
Comparison of thigh tubercles; T. graeca vs. T. whitei
There is a considerable difference in the construction of the thigh
tubercles of the two species. Those of specimens conforming to the
holotype of T. graeca L. are smaller, and typically do not extend
beyond 2.5mm from the surface. Those of T. whitei BENNETT are whitish
in colour and typically extend in an inwardly curved direction (towards
the cloaca) as much as 7mm from the surface of the skin, having sharply
pointed ends. This form of thigh tubercle has not been observed in any
specimen conforming to the Linnaean holotype, but is consistently
found in T. whitei BENNETT (see plates 7 and 8).
Original publication
In the 1836 edition of Whites "The Natural History of Selborne" the
editor, Bennett, added the following footnote to letter L of 1781
concerning "the old Sussex tortoise" (p.360 - 361);
"Interesting as the old family tortoise has been rendered by the
anecdotes related of him by Gilbert White, his history may be closed by
the statement that his life was not prolonged much beyond that of his
protector. He died, it is believed, in the spring of 1794; after an
existence extended in England to about fifty-four years, the last
fourteen of which were spent at Selborne. The thick shell, in which he
was coffined while alive, is preserved in the residence of the master
who secured for him an enduring existence in the memories of many.
My friend Mr. Bell regards the specimen, which he has had an
opportunity of inspecting, as an old and worn shell of the bordered
tortoise, Test. marginata, SCHOEPFF: and all who are acquainted with the
extent and accuracy of his knowledge of the Testudinata, must be aware
that anyone who differs from him on such a subject, is probably in the
wrong. Yet at this risk I have ventured to regard the Selborne tortoise
as a distinct species. Its shell is less elevated than is usual in the
bordered tortoise, once named on that account the bell-shaped: its
wrinkles are less strongly marked and less sharp: its sub-caudal plates
form with each other a much more open angle: and its anterior
supra-femoral plate, instead of running to a point towards the back,
has an inner margin nearly of equal length with its anterior and
posterior edges. But the general form of the shell of a tortoise, the
sculpture of its surface, and the shape of particular plates, are all
too variable in many species to warrant the adoption of any or all of
these characters as absolutely distinctive; and on them no assured
reliance can consequently be placed. More stress may be laid on the
animal, and on particular organs or plates attached to its body; and in
the case of Gilbert White's tortoise there is a fragment remaining of
the skin of one of the thighs which principally induces me to regard it
as distinct from the bordered species: for on this fragment of skin
there is a large white conical process or spur. No such process was
noted by Mr. Bell on the specimen of the bordered tortoise which he had
alive and which is beautifully figured in his splendid work on the
Testudinata: evidence, it is true, of a negative character only, but
becoming positive when taken in conjunction with the distinct statement
of M. Bibron, (in the Erpetologie General, which he is now publishing
in conjunction with M. Dumeril,) that there are no large horny
tubercles in that species on the hinder part of the thighs. Although
the bordered tortoise is far from uncommon in Greece, and in other
countries on the shores of the Mediterranean basin, I am compelled to
refer to authorities for its structure, as I am not aware of the
existence in london of a living or preserved specimen of the animal.
Mr. Bell has the only two shells of it that are known to me.Mrs.
White's, for the loan of which I am indebted to her kindness, may be a
third: but it seems to me, with our present knowledge on the subject,
that it must be regarded as distinct.
I propose for it the name of Testudo whitei."
As Bennett states in his footnote, at this time the opinion of Bell
was that Whites tortoise represented a specimen of Testudo marginata
SCHOEPFF 1792. A view which was obviously still held by J. E. Gray as
late as 1870, for in his "Supplement to the Catalogue of Shield
Reptiles" (p.11) he writes of it;
Female, or var. Whitei
Testudo whitei, Bennett in White's Selborne.
"A fine adult shell, with the hinder margin moderately expanded, and the
caudal shield bent down and slightly inflexed; the sternum flat, the
hinder lobe tapering behind, the anal shields being not more than
two-thirds the width of the shields before them,; the pectoral shields
very short, not above one-fourth the length of the abdominal shields on
the inner half, the outer half about double the width and square; the
dorsal shields black; the areola varied with yellow; the hinder upper
half of the marginal shields varied with yellow; the underside is
yellowish white, with a few irregular unequal-sized black spots. Adult
female. The tortoise was described in White's 'Selborne', and presented
to the Museum in 1858 by Mrs. Christopher, niece of Mr. White."
Gray obviously regarded the White specimen as somewhat unusual, for it
is referred to as Testudo marginata var. whitei. In fact, Bennett was
entirely correct in his original dismissal of it as a specimen of
marginata. Identical errors on the parts of authorities of the
undoubted taxonomic abilities of Gray and Bell do, however, serve to
demonstrate that these tortoises definitely have been mistaken for
marginata - a fact which goes some way to explaining the large number
of otherwise inexplicable reports of Testudo marginata encountered in
much 19th C. literature on the chelonian fauna of Algeria.
A somewhat puzzling feature of Bennetts description is his failure to
refer (in any way) to Linnaeus's Testudo graeca, Testudo pusilla or
Testudo tessellata minor africana (Edwards, 1748). Confronted with an
enigmatic tortoise, it seems odd that none of these other possibilities
was even discussed - particularly as White himself had ventured the
opinion that his tortoise was a T. graeca. The probable answer is that
Bennett recognised immediately that Whites tortoise was so different
from Testudo graeca L. that he did not feel that any question of
confusion arose, so he simply did not feel it necessary to mention them
or draw comparisons with them. It is also noteworthy that such eminent
authorities as Gray and Bell also did not connect this specimen with
the Testudo graeca of Linnaeus. The dimensions of the specimens must
have suggested that the only really likely candidate was Testudo
marginata - an animal which does indeed closely approximate Testudo
whitei in size, whereas T. graeca L. manifestly does not.
Conclusions;
Testudo whitei BENNETT 1836 is not as has often been thought a
synonym of Testudo graeca L. 1758, but is a full and independent
species. Morphological differences between the two races are constant,
and T. graeca L. never attain the dimensions reached by T. whitei
BENNETT. Early reports of North African tortoises which approach the
size of T. marginata SCHOEPFF 1792 may be found to concern T. whitei
BENNETT. To date, bibliographic research has revealed no earlier
credible description of this species than that published by Bennett in
1836, so his nomenclature Testudo whitei should be regarded as valid.
No type locality was specified in Bennett's original publication, but
the terra typica of this species is undoubtedly Algiers and its
environs, Algeria.
Some synonyms in lit.:
As Testudo marginata SCHOEPFF; Gervais 1836; Gray 1870; Lallemant 1867.
As Testudo ibera PALLAS: Boulenger 1889; Lortet 1888; Johnson 1928.
As Testudo graeca graeca; Flower 1945; Lambert 1982; Loveridge and
Williams 1957; Pritchard 1966 and 1979;
Authors note;
The re-discovery of this tortoise changes the entire position regarding
the status and nomenclature of the North African members of the genus
Testudo. It also raises a number of vitally important questions
concerning current population status and conservation. Hitherto, data
from T. graeca L. and T. Whitei have not been considered separately. As
a result, figures for T. graeca L. are distorted, and figures for T.
whitei non-existent. We know next to nothing of the ecology and
behaviour in the wild of T. whitei, and have no data whatsoever
regarding the current population status of the species. Few surveys of
the herpetofauna of Algeria have been published in recent years, and of
those that have the picture in respect of the tortoises is not
encouraging (Sura, 1987). We do know that the species was heavily
exploited commercially (Lambert, 1980) for many years to supply the
pet trade in Europe (large numbers being collected for export to
France, Germany and the U.K prior to CITES and other export
restrictions), but we have no recent data on distribution or numbers.
This information is required urgently to assess if further conservation
steps need to be implemented in order to secure the species continued
survival in its natural habitat.
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