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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 AA INSTITUTO 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,