An Inventory ofCombat Rituals in Snakes
CHARLES C. CARPENTERDepartment of ZoologyUniversity of Oklahoma
SMITHSONIANHERPETOLOGICAL INFORMATIONSERVICENO. 691986
SMITHSONIANHERPETOLOGICALINFORMATIONSERVICE
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One interpretation of the medical caduces symbol fromantiquity is that it represents the lyre-shaped configurationof two snakes contesting one-another. Such a belief may havebeen derived from observing the combat ritual of Elaphelonqissima which assumes a lyre-like posture during this typeof interaction and is called the Aesculapian Snake after theGod of Medicine. If this interpretation is correct then thepresence of this combative behavior in snakes has beenrecognized since ancient times.However, it is not until the early part of this centurythat references to combat rituals in snakes began to appear inthe scientific literature, and in recent years the list ofreferences has grown to now include a breadth of species inthe families Boidae, Colubridae, Elapidae and Viperidae(Viperinae and Crotalinae).It is the purpose of this inventory to bring together alist of the species of snakes for which, in my interpretation,the phenomenon of combat ritual has been authenticated viapublished literature sources. I also hope that this inventorywill stimulate others to watch for this type of behavior,refer to some of the most recent descriptions for methods ofobserving and describing, and help to lengthen this list. Inthis way we will all contribute to grasping a betterunderstanding of what snake combat rituals mean and how theymay have evolved.
FAMILY ANDGENUS SPECIES CITATIONBOIDAECorallusLiasisM orel iaPythonSanziniaCOLUBRIDAE-Coluber
Coronella
DiadophisDrymarchon
caninachildrenispilotes
molurusmadaqascariensis
constrictorqemonensis
viridif lavusaustriaca
punctatuscorais
Osborne, 1984Ross, 1973Ross, 1978Covacevich, 1975Shine, et al. , 1981Barker, et al., 1979Carpenter, et al., 1978Shine, et al., 1981
Bogert and Roth, 1966Shaw, 1951Davis, 1936Shaw, 1951Stemmler-Morath, 1935Guibe and Saint Girons,1955Andren and Nilson, 1976Bogert and Roth, 1966Davis, 1936Fitch, 1975Rollinat, 1947Shine, 1978Smith, 1943Smith, 1954Fitch, 1975Bogert and Roth, 1966Murphy, 1973Shine, et al., 1981Tinkle, 1951Waide and Thomas, 1984
Elaphe guttatalonqissim a
obsoleta
quadrivirqataLampropeltis calligastergetulus
mexicanapyromelanatrianqulum
Malpolon monspessulanusMasticophis f lagellumtaeniatus
Murphy, 1973Shaw, 1951Bellairs, 1970Carpenter and Ferguson,1978Murphy, 1973Pope, 193 7Shaw, 1951Shine, et al., 1981Stemmler-Morath, 1935Brecke, et al., 1977Gillingham, 1980Littleford and Keller, 1946Mitchell, 1981Rigley, 1971Shine, 1978Shine, et al., 1981Stickel, et al., 1980Fukada, 1961Shine, 1978Carpenter, 1984Moehn, 1967Shine, et al, 1981Carpenter, 1977Carpenter and Gillingham,1977Clark, et al., 1984Lewke, 1979Neitman, 1980Murphy, et al., 1978Shine, et al., 1981Martin, 1976Fitch and Fleet, 1970Murphy, 197 3Shaw, 1951Shine, et al., 1978Shaw, 1951Kennedy, 1965Bennion and Parker, 1976Parker, 1974Shine, 1978Shine, et al . , 1981
Pituophis melanoleucus
Pseudaspis canaPtyas (Zamenis) mucosus
Rhinocheilus leconteiSonora episcopa (semianulata)
Theltornis . capensisELAPIDAEAustralelaps superbusBunqarus
CryptophisDemansia
candidus
Bogert and Roth, 1966Carpenter and Ferguson1978Lowe, 1948Murphy, 1973Parker, 1974, 1980Shaw, 1951Shaw and CampbellShine, 1978Shine, et al, 1981Woodbury, 1941 1974FitzSimons,Shine, 197E 1962Abdulali, 1941Beadon, 1910Bhattacharya and Bhatnager,1968Davis, 1936Lowe, 1948McCann, 1935Millett, 1909Pope, 1937Prater, 1933Shaw, 1951Shine, 1978Wall, 1921Osborne, 1984Kassing, 1961Kroll, 1971Shaw and Campbell, 1974Shine, 1978Shine, et al., 1981Bennefield, 1982Shine and Allen, 1980Shine, et al., 1981Shine, 1978Taylor, 1965Wall, 1921Whitaker, 1971Shine and Allen, 1980Shine, 1980Shine and Allen, 1980
Dendroaspis
HemiaspisNaia
jamesonipolylepis
dameliihaje
Bogert and Roth, 1966Grant, 1956Leloup, 1964Bogert and Roth, 1966Leloup, 1964Shine, 1978Carpenter, and Ferguson,1978Broadley and Cock, 1975Leloup, 1964Shine, 1978Shine and Allen, 1980Richards, 1885FitzSimons, 1962Shine, 1978
tripudians
Bogert and Roth, 1966Pope, 1935Shine 1978Thomas, 1970Wall, 1921Deoras, 1971Kopstein, 1914Shine, 1978Notechis scutatusOphiophagus hannahOxyuranus scutellatus Shine, 1977, 1978Oliver, 1956Hosmer, 1953Shine and Allen, 1980Shine, et al. , 1981Worrell, 1963Pseudechis porphyriacus
Pseudona ja texti li s( =Deinansia
)
Baker, 1968Carpenter and Ferguson,1978Fleay, 1937a, 1951Lowe, 1948Shaw, 1951Shine, 1977, 1978Shine, et al, 1981Worrell, 1963Carpenter, and Ferguson,1978Fleay, 1937a, 1937b, 1951Shaw, 1951Shine, 1978Shine and Allen, 1980Worrell, 1963
VIPERIDAEVIPERINAEBitis arietans
Causus
caudalisqabonicadef ilippilichtensteiniiresimus
Bogert and Roth, 1966Carpenter, 1977Harvey, 1964Pitman, 1974Rose, 1962Shine, 1978Thomas, 1959, 1960, 1972Akester, 1983Akester, 1979a, 1979bBogert and Roth, 1966Akester, 1979aLoveridge, 1942Schmidt and Inger, 1957Bogert and Roth, 1966Curry-Lindahl, 1956
rhombeatusVipera
ammodytes
Broadley and Cock, 1975Bogert and Roth, 1966Davis, 1936Froesch-Franzon, 1971Noble, 1937Pope, 1937Prior, 1933Carpenter, 1977Laue, 1974Shine, 1978Steward, 1971Thomas, 1959, 1960, 19611969, 1971aspis Baumann, 1929Bogert and Roth, 1966Duguy, 1958Guibe and Saint Girons1955Koster, 1932Naulleau, 1967Reuss, 1926Saint Girons, 1949Shine, 1978Steward, 1971
berus Andren, 1975, 1976, 1981Baumann, 1929Bernstrom, 1958Bogert and Roth, 1966Carpenter and Ferguson,1978Davis, 1936Hajek-Halke, 1941Heilborn, 1930Kellaway, 1982Lotherington, 1934Nilson and Andren, 1982Prestt, 1971Prior, 1933Reuss, 1923, 1926Saint-Girons, 1952Schmidt and Inger, 1957Shine, 1978Smith, 1954Thomas, 1955, 1959, 1960Viitanen, 1967Volsoe, 1944lebatina Carpenter, 1984Stemmler, 1967macrops Kopstein, 1914russelli Laue, 1974CROTALINAEAgkistrodon Bogert and Roth, 1966Carpenter and Gillingham,In pressFitch, 1960Gloyd, 1947Noble, 1937Ramsey, 1948bilineatus Burchfield, 1982Carpenter and Gillingham,In press
contortrix Bogert and Roth, 1966Carpenter and Gillingham,In pressFitch, 1950Gloyd, 1947Life, 1950Mitchell, 1981Shaw, 1948, 1951Shine, 1978Shine, et al., 1981Stewart, 1984Sutherland, 1958
Bothrops qodmanimoojeni
Allen and Swindell, 1948Bogert and Roth, 1966Burkett, 1966Carpenter, 1977Carpenter and Ferguson,1978Carpenter, et al., 1976Carpenter and Gillingham,In pressCarr, 1963Carr and Carr,Davis, 1936Ellis, 1979Martin, 1984Perry, 1978Ramsey, 1948Shine, 1978Shine, et al.
,
Wharton, 1966,
1942
19811969Wright and Wright, 1957Carpenter, 1984Leloup, 1975Calloselasma rhodostoma York, 1984Carpenter and Gillingham,In PressCrotalus Arnold, 1935Bogert and Roth, 1966Carpenter, 1984Crites, 1952Gloyd, 1947, 1948Klauber, 1956Lowe, 194 2Lowe and Norris, 1950Shaw, 1948Shine, et al., 1981Sutherland, 1958Wagner, 1962Wright and Wright, 1957
adamanteus Bogert and Roth, 1966Klauber, 1956Shine, 1978Wagner, 1962atrox Bellairs, 1970Carpenter, 1977, 1984Carpenter and Ferguson,1978Foree, 1949Gillingham, et al., 1983Gloyd, 1948Grimpe, 1984Jenni, 1966Kennedy, 1942Klauber, 1956Lowe, 1942Murphy, 1973Shaw and Campbell, 1974Shine, 1978Wagner, 1962Whisenhunt, 1949basiliscus Davis, 1936cerastes Bogert and Roth, 1966Klauber, 1956Lowe and Norris, 1950Shaw and Campbell, 1974Shine, 1978Shine, et al., 1981durissus de Langlada, 1975Shine, 1978horridus Bogert and Roth, 1966Klauber, 1956Murphy, 1973Shine, 1978Shine, et al, 1981Sutherland, 1958lepidus Carpenter, 1977Carpenter, et al., 1976Shine, 1978mitcheli Bogert and Roth, 1966Klauber, 1956Shaw and Campbell, 1974Shine, 1978Sutherland, 1958
10ruber Carpenter and Ferguson,1978Eibl-Eibesfeldt, 1961Evans, 1955Hediger, 1955Klauber, 1956Pequegnat , 1945Shaw, 1948Shaw and Campbell; 1974Shine, 1978Shine, et al. , 1981scutulatus Klauber, 1956Shine, 1978Shaw and Campbell, 1974Carpenter, 1984viridis Bogert and Roth, 1966Bromley, 1934Carpenter, 1977, 1984Dayton, 194 9Gehlbach, 1965Gloyd, 1947Klauber, 1956Shaw, 1948Shine, 1978Shine, et al., 1981Staley, 1983Sutherland, 1958Thorne, 1977Sistrurus miliarius Carpenter, 1977, 1979Lindsey, 1979Palmer and Williamson, 1971Shine, 1978Shine, et al., 1981
11LITERATURE CITED
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Baker, A. B. 1968. Snakes in combat. Proc. Roy. Soc. New SouthWales 1966-67:29-31.Barker, D. G., J. B. Murphy and K. W. Smith. 1979. Socialbehavior in a captive group of Indian pythons. Pythonmolurus (Serpentia, Boidae) with formation of a linearhierarchy. Copeia 1979:466-471.Baumann, F. 1929. Experimente uber den Geruchssinn und denBeuteenwerb der Viper ( Vipera aspis L.) . Z. Vgl. Psysiol.10:36-119.Beadon, C. 1910. A snake flirtation. J. Bombay Natur. Hist.Soc. 20:288.Bellairs, A. 1970. The Life of Reptiles. Universe Books,New York (2 vols )
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12
Brecke, B., J. B. Murphy and W. Seifert. 1977. An inventory ofreproduction and social behavior in captive Baird's ratsnakes. Herpetologica 32:389-394.Broadley, D. G. and E. V. Cock. 1975. Snakes of Rhodesia.Longman Rhodesia, Ltd.; Salisbury.Bromley, R. 1934. Rattlesnake combat. Nature. Mag. 24:45.Burchfield, P. M. 1982. Additions to the natural history of thecrotaline snake Aqkistrodon piscivorus (Reptil'ia). J.Herpetol. 16:376-382.Burkett, D. R. 1966. Natural history of cottonmouth moccasin,Aqkistrodon piscivorus (Reptilia). Univ. Kansas Publ.Mus. Natur. Hist. 17:435-491.Carpenter, C. C. 1977. Communication and displays of snakes.Amer. Zool. 17:217-224.Carpenter, C. C. 1979. A combat ritual between two male pygmyrattlesnakes ( Sistrurus miliarius ). Copeia 1979:638-642.Carpenter, C. C. 1984. Dominance in snakes, pp. 195-202. IjnR. A'siegel, L. E. Hunt, J. L. Knight, L. Malaret. N. L.Zuschlag (eds.) Vertebrate Ecology and Systematics - ATribute to Henry S. Fitch. Univ. Kansas Mus. Nat. Hist.Spec. Publ. 10.Carpenter, C. C. and G. W. Ferguson. 1978. Variation andevolution of stereotyped behavior in reptiles, pp. 335-554.In C. Cans and D. W. Tinkle (eds.) Biology of theReptilia. Vol. 7. Academic Press, New York.Carpenter, C. C. and J.C. Gillingham. 1977. A combat ritualbetween two male speckled kingsnakes ( Lampropeltis qetulusholbrooki : Colubridae, Serpentes) with indications ofdominance. Southwest. Natur. 22:517-524.Carpenter, C. C. and J. C. Gillingham. 1985. (In Press)Ritualized behavior in Aqkistrodon and allied genera. InR. Conant and H. K.Gloyd (eds.) Monograph on Aqkistrodonand Allied Genera. Soc. Study Amphib. Rept. Spec. Publ.Carpenter, C. C, J. C. Gillingham and J.B. Murphy. 1976. Thecombat ritual of the rock rattlesnake ( Crotalus lepidus ).Copeia 1976:764-780.Carpenter, C. C, J. B. Murphy and L. A. Mitchell. 1978. Combatbouts with spur use in the Madagascan boa ( Sanziniamadaqascariensis . Herpetologica 34:207-212.Carr, A. 1963. The Reptiles. Time Inc.; New York.Carr, A. amd M. H. Carr. 1942. Notes on the courtship of thecottonmouth moccasin. Proc. New England Zool. Club. 20:1-6.Clark, D. L., J. C. Gillingham and A. Rebischke. 1984. Notes onthe combat behavior of the California kingsnake,Lampropeltis qetulus californiae , in captivity. BritishJ. Herpetol. 6:380-382.Covacevich, J. 1975. Snakes in combat. Victorian Natur.92:252-253.Crites, A. S. 1952. A Hunter's Tale of the Great Outdoors.Ward Ritchie Press; Los Anqeles.Curry-Lindahl , K. 1956. Mambas in combat. African Wildl.10:340-341.Davis, D. D. 1936. Courtship and mating behavior in snakes.Field Mus. Natur. Hist. Zool. Ser. 20:257-290.
13
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14
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15
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