INTRODUCTION

Cephalopods may be divided into five types according to their buoyancy. Members of several families such as the Octopodidae, Loliginidae and Ommastrephidae are negatively buoyant and must swim to stay in midwater and are therefore highly muscular animals. Others have mechanisms to make them neutrally buoyant so they can remain suspended in midwater without effort. Nautilus, Spirula and cuttlefishes have low pressure gas-filled chambers and their flesh is muscular and non-buoyant (Denton & Gilpin-Brown, 1973). Squids of one family, the Gonatidae, have a low density oil in their livers to give buoyancy but most of their body is muscular. Some oceanic octopods have very watery tissues in which lighter chloride ions replace sulphate ions (Denton & Shaw, 1961). In 12 of the 26 teuthoid families the buoyancy is provided by low-density ammonia-rich solution in their body and head tissues or in an expanded coelomic cavity (Clarke, Denton & Gilpin-Brown, 1979). These ammoniacal squids are extremely abundant in the oceans of the world and form a large part of the diet of birds, cetaceans, seals and fish (Clarke, 1977). When their biomass is estimated from their utilization by predators it is important to know their properties as food and, in particular, their calorific values. As pointed out by Croxall & Prince in a review of the calorific values of cephalopods (1982), all the known values are of muscular, negatively buoyant species because they are of value as food for humans but no measurements have been made on the ammoniacal or oily species which are probably as important, or even more important, in the economy of the ocean (Clarke, 1983).