Observations on mating in Mediterranean Sepiola and Sepietta species and review of mating behaviour in Sepiolinae (Cephalopoda: Sepiolidae)

Abstract Mating was observed and described in captive individuals of Sepiola affinis, Sepiola intermedia and Sepietta obscura (Cephalopoda: Sepiolidae) collected in the Catalan Sea, western Mediterranean Sea. This is the first report of a mating event in S. intermedia; it is also the first detailed description of the mating behaviour for the other two species. The published literature on mating in Sepiolinae, which includes both cursory reports and in-depth descriptions of mating events, was thoroughly reviewed. In all, copulation has been examined in eight species belonging to four different genera, namely, Eumandya, Euprymna, Sepietta and Sepiola, starting from 1894 to the present. Common traits of the mating behaviour were detected among the studied sepioline species, so that a general five stages succession of actions is established to portray the mating progress in Sepiolinae: (A) female hovers by, male attention (it is discussed whether actual copulation is preceded by any courtship); (B) male approaches female from below; (C) male grasps female at the neck by its third arms, inserts its first arms in the female’s mantle cavity (the hectocotylised left arm is thus aligned with the bursa copulatrix), holds the female’s mantle by its second arms and positions itself and mate in the “parallel position”; (D) copulation and transfer of spermatophores from male to female (this stage may last from 3 min to 3 h); (E) mating dissolution. Mating occurs preferentially during the dark hours; it is described as violent and the female tries to escape the forceful grasp by the male; the male skin coloration turns darker. The similarity of the mating behaviour in all examined sepioline species is an evidence of both its evolution in harmony with their copulatory organs (hectocotylus and bursa copulatrix) and, seemingly, its common derivation to the whole Sepiolinae clade