XX (ZS700U_ 



Bull. nut. Hist. Mas. Loud. (Zool.) 68(2): 91-99 



Issued 28 November 2002 



Atractaspis (Serpentes, Atractaspididae) the 

 burrowing asp; a multidisciplinary minireview 



E. KOCHVA 



Department of Zoology, Tel Aviv University, Ramat Aviv, 69978 Tel Aviv, Israel 



CONTENTS 



Dedication 91 



History 91 



Taxonomy 91 



Distribution 92 



Behaviour 92 



Venom apparatus 93 



Venom 93 



Evolution 97 



Acknowledgements 98 



References 98 



SYNOPSIS. The family Atractaspididae is a highly modified derivative of a lineage that apparently arose early in the history of 

 'colubroid' snakes, and its taxonomy and relationship with other ophidian groups is still uncertain. Snakes of the genus 

 Atractaspis have a characteristic venom apparatus, including the structure and function of the striking unit and of the venom 

 glands. The composition of their venom is also unique in containing several low-molecular weight components, thesarafotoxins, 

 which affect the cardiovascular system and are similar to the mammalian endothelins. 



Dedication 



This paper is dedicated to Dr. Garth Underwood on the occasion of 

 the 35 ,h anniversary of his classic 'Contribution to the Classification 

 of Snakes' ( 1967), about which one may say: 



This is a small book by a great man! 



And also -(T ,il ,T"Q) /7J77Z7/7 77A 7 OJJ7H pVUHOJ ItiJT] 

 A little that contains a lot (Theodor & Albeck, 1996) 



HISTORY 



There are not many snake species that posed problems from the very 

 beginning of their discovery; one of the most prominent ones is 

 certainly that which we now call the genus Atractaspis of the family 

 Atractaspididae (Fig. 1 ). 



The first two specimens of Atractaspis were described by Reinhardt 

 in 1843 as Elaps irregularis, a species that he considered to be 

 extremely abnormal because of the presence of only a few, very 

 small teeth. On the basis of squamation, Underwood inferred that at 

 least one of the specimens was A. dahomeyensis. The genus 

 Atractaspis was established by Smith in 1 848 for the South African 

 species bibroni and since then it was variously considered as a 

 separate family, as a subfamily of the family Viperidae, and finally 

 as a genus within the Viperinae. 



Already Haas (1931), on the basis of the pattern of the head 

 musculature, was unhappy with the inclusion of Atractaspis in the 

 Viperidae, but it was not until 1961 that Monique Bourgeois, 

 studying at the Universite Officielle du Congo a Lubumbashi, came 

 out with a challenging question: Atractaspis - a misfit among the 



Viperidae? This short note was followed by a detailed study suggest- 

 ing the establishment of a separate subfamily for a group of 

 opisthoglyphous colubrids together with Atractaspis (Bourgeois, 

 1968). 



Underwood (1967) lists a long series of skeletal and other ana- 

 tomical characters in which Atractaspis differs from the Viperidae 

 and states: '■Atractaspis differs so widely from the other vipers that 

 I have no doubts about reviving a separate family group taxon to 

 receive it' (p. 103). This he did after a detailed analysis that resulted 

 in the resurrection of the subfamily Atractaspidinae (Gunther, 1 858); 

 and, finally, the establishment of a separate family. Atractaspididae, 

 for the approximately 15 species of Atractaspis together with some 

 African colubrid genera (Underwood & Kochva, 1993). 



TAXONOMY 



Recently several suggestions concerning the taxonomy and relation- 

 ship of the Atractaspis species have been raised, mainly dealing with 

 the question of which additional genera should be included in the 

 family Atractaspididae and with which, if any, larger clades they 

 should be grouped (Gravlund, 200 1 ). Underwood himself (personal 

 communication) is now reconsidering the composition of the family 

 Atractaspididae in order to decide which genera, in addition to 

 Atractaspis, should be included in it. However, no one is now 

 questioning the separate status of the genus Atractaspis, and its 

 apparent distinction from the other venomous snake families is 

 widely, though not unanimously, agreed upon. Atractaspis thus 

 certainly deserves the rank of a family of its own; this may also 

 include some rear-fanged snakes that are apparently harmless as far 

 as humans are concerned. 



> The Natural History Museum. 2002 



