Ie Sys
Pe Re!
ZOOLOGISCHE MEDEDELINGEN
UTGEGEVEN DOOR HET
RIJKSMUSEUM VAN NATUURLIJKE HISTORIE JE LEIDEN
(UNISTERIE VAN CULTUUR, RECRRATIE EN MAATSCHAPPELIJK WERK)
Deel 41 no. 11 27 juli 1966
‘THE PATTERN OF DERMAL-VERTEBRAL
CORRELATION IN SNAKES AND AMPHISBAENIANG
by
A. ALLAN ALEXANDER?) and CARL GANS:
Pepartment of Biology, State University of Now York at Buffalo,
Bullalo, N. ¥. 4214, U.S.A.
Irropuetion
It has long been known that the arrangement of external scales retains a
constant relation to the priviary pattern of mesodermal segmicntation, The
ratio of the number of dermal scale rows or annuli to the number of verte
brae has, therefore, been considered io be of fundamental importance im
squamate chssification (Stehfi, 110; Camp, 1923). Yer the diffieulyy of
determining i( has fostered the presentation of hypotheses tased on relatively
fevr total counts; some suck hypotheses have relied pom comparisons made
by dissection along limited portions of the trunk, The general implication
communicated by the many statements in the Hterature has been that the
ratias are constant at the generic or familial level, that the ratios ordinarily
represent simple, whole number relations (ie 1 if, £2, 2:2), and that
the scale-vertebra relation is constant along the Yength of the body
Authors have disagreed regarding the presumed evolution of these rei
larities; thus Stebli (1910) argued that the 1: 1 ratio was primitive, while
Camp (1923) considered it to be most highly advanced, The most “primitive”
snakes (Bellairs & Underwood, 1951) were generally stated to have a 2 51
and the “fadvaneed” forms ar: 1 ratio, In contrast, the presumably most
primitive amphisbaenid (Smalian, 1884) has generally been stated to have
at: tand all other, and prestumably more advanced, forms, a 2: 1 ratio.
A recent report (Gans & Taub, 1963) showed a simple method of deter
‘mining these ratios, emphasized that the ratios varied markedly at the species
1) Pretent address: Department of Biology,
sys College, Buffalo, NY. 14208,172 ZOOLOKISCHE MEDEDELINGEN $1 (4y66)
level, and also showed that the relationship of scales to vertebrae within
the ‘Typhiopidae was not a simple, whole nursber ratio. This suggested the
desirability of checking the actual valves om which the varioss rales had
been established.
We are grateful to Drs, Neil D, Richmond and 1, W, Parker for volun
tering information on aspects of this problem, and to Ds, K, I Wiliams
for reading the manuscript. The specimens examined were loaned by C.
M. Bogert and Ik. G. Zuveifel of the American Museum of Natural History,
New York (AMNED}, A. E, Leviton of the California Academy of Sci
(GAS), the Carl Gans collection at Buffalo (CG), N. D. Richmond of the
Carnegie Museum, Pittsburg (CM), H. Marx: and RoE. Inger of the
Chicago Natural History Museum (CNUIM), A. R, Hoge of the Institute
Butattian, Sao Paulo (IB), P. W. Braestrup of the Universitetets Zoologishe
“Museu it Copenhagen (KM), J. AON. Cranwell and J. M. Gallardo of
the Muse Argentina de Ciencias Naturales in Buenos Aires UMACN),
EE. Williams of the Muscunt of Comparative Zoology (MCZ), M. Boese=
man of the Rijksmusenm van Natuurlifke 1 en (RMNHD. H.
Wermuth of the Staatlichen Museum fir Naturkunde, Seutygart (SMNS),
C.F. Walker of the University of Michigan Museum of Zoology (UMMZ),
1D. M. Cochran of the United States National Museum (USNM}, and J
Elselt of the Naturhistorisches Museum, Vienna (VM). Dr. J. C. List
kindly percnitted us to cite data from his (unpublished) thesis, This study
is supported by a grant from the National Science Foundation GR-2960
(to C. Gans),
Mernops
The arrangement and numbers of scale rows, or annuli, on the dorsal
and ventral surface of trunk and tail, and the numbers of trunk and caudal
vertebrae were determined for some 220 specimens, Similar determinations
were accomplished on 130 specimens of Banus, Additionally, determinations
fom some 100 other specimens recorded by Gans & Taub (1965), Gans &
Laurent (1965), and by List (1956, unpublished thesis) were also used in
the analysis, The ranges of observed values are shown by species in table 1
for the amphisbaenidls, and in table 2 for the “primitive” snakes and # few
advanced snakes,
The forms sampled include the 23 recognized genera of amphishaentds,
fifteen genera of primitive snakes, and various advanceal snakes, ‘The various
ratios observed in each family or other grouping ace shown in table 3.
‘The numbers and proportions of vertehrae were determine for X-rays
taken with an unfiltered 60 KVP dental unit at 3. ma on Kodak Type AAINSTITUTO
OSWALDO CRUZ
BLOT. <:.,
' industrial film; the target to film distance was 50 inches. All determinations
were made uuder a dissecting microscope, Measurement of proportions ab-
viously had to be restricted to those vertebrae X-rayed while lying with their
long axis in parallel to the film, Hypague and fine steel pins were used
to indicate the position of dermal segments relative to skeletal structures
Checks inticated that the counts of dermal and vertebral elements along
the trunk were repeatable with less than 19% error, Caudal structures, parti=
ularly in short-tailed forms, are markedly reduced in size and the vertebrae
are often partially fused, The differences hetween repeated counts here ap-
{Hostal fore” Corton rma dea dene to 1
poses a problem, partictlary in the typhlopids, Hese the tal is shor, sharply
curved, and composed of smaller vertebral elements. Additionally the X-rays
i suggest that the soft tissues may shift (possibly post-mortem) 0 that the
cloacal slit may, im a given species, fall anywhere from the third caudal to
the fourth from last trunk vertebra. Tt may be that this accounts for some
ai the apparently abersant ratios; otherwise it may be that spevies differences
are being dealt with,
ALEXANDER & CANS, DERMAL-VERTERRAL CORRELATION
Resuicts
Serpeates
Tronk proportions, -~ Neither the Tengih of the dermal segments nor of
the vertebrae remains constant throughout the «unk region, ‘The changes
ase gradual, but quite evident when comparisons are made between regions.
Both seales and vertebrae of the muchal-pectoral and precteacal regions
ave generally of equivalent length, and defintely shorter than those of the
imidthoratic region. ‘The lengths of the scales and vertebrae generally change
in parallel, maintaining a constant eclationship along the trunk, and giving
rie to the desczibed simple, whole number ratios (je, 1: t, 21 1, etc.). This,
{s im accord with the expected hasie embryologicai relationships of the dermal
and vertebral clerents. In a few foros, the coustancy of the relaionship is
disrupted hy (seemingly sevondary) changes in the dermal elements, changes
that result in deviant ratios
Richmond (in litt.) commented on the marked regional differences in
scale size seen in certain typhlopfds. Vet ehe vertebral conn of these forms
shows the same regioral proportions describe! above for other snakes, ex-
cept perkaps for a shorter precloacal region, and a sharp change in the length
of vertebrae between the thoracic and precloacal zanes. The dermal elements
also increase in length along the trunk, but seemingly with no corresponding
dlecrease in the short precioacal region. The degree of increase appears to
be species specific,