NAFO SCI. Coun. Studies, 9: 125-132 Predatory R of the Flying Squid (Todarodes sagittatus) in North Norwegian Wale Anne Breiby and Malcolm Jobling Institute of Fisheries, University of Troms¢ N-9001 Troms¢, Norway Abstract Analysis of the gut contents of Todarodes sagittatus were carried out from samples which were obtained by jigging in coastal waters of northern Norway. The food was comprised predominantly of pelagic species, with polychaetes, crustaceans and fish being the major prey groups. Feeding indices provided evidence of a size-related shift in dietary composition, with increased reliance on fish prey as the squid increase in size. Use of otolith size-fish size relationships to estimate prey size indicated that the range of prey size increased with squid growth. The presence of relatively undigested pieces of squid in the gut contents was interpreted as being an artefact of the fishing technique, and it is suggested that the importance of cannibalism may have been overevaluated in other studies on the feeding habits of squids. The ways in which choice of sampling and analytical techniques may influence the results of dietary studies are discussed. I ntrodl.lction Materials and Methods Although the European flying squid (Todarodes Samples of T. sagittatus were collected at various sagittatus) is thought to occupy a key position in the times from October 1982 to October 1983. Although it food web of Arctic regions in the Northeast Atlantic, would have been desirable to have collected the sam­ comparatively little is known about its feeding biology, ples at one specific location, movements and availabil­ with the majority of available information being widely­ ity of squid made this impossible, Consequently, the scattered, unsystematic lists of the types of food orga­ samples were taken at several locations along the nisms (Wiborg, 1972, 1978,1979,1980,1981; Jonsson, north coast of Norway between Andenes and Kam(by­ 1980). The feeding habits of squids from other regions fjord (Fig, 1). Squid were captured by jigging in depths have been examined in more detail and the results of from the surface to 50 m. Jigging was not limited to several studies have shown that they feed on pelagic or particular periods of the day, and most of the samples semi-pelagic prey (Squires, 1957; Vinogradov and include squid which were obtained at different times Noskov, 1979; Macy, 1982; Mangold 1983; O'Sullivan within a given 24-hr period, Ten samples, varying in and Cullen, 1983). Although some authors have size from 10 to 170 individuals, were collected during reported that there is a change in dietary composition the course of nine research cruises. Relatively few (97) with increasing size of squid (see review by Mangold, male squid were captured, and the numbers in the 1983), most of these studies have been descriptive in samples varied between 0 and 53 individuals. Conse­ nature and few attempts at detailed analysis have been quently, this study refers predominantly to examina­ made, tion of the feeding habits of female squid, In the present study, the feeding habits of T. sagit­ Immediately after capture, the dorsal mantle tatus have been investigated and possible mechanisms length (ML) of each squid was measured to the nearest which lead to dietary changes are examined. Attempts 0.5 cm, and the alimentary tract was usually removed, were made to compare the findings with results of frozen in liquid nitrogen and stored at -200 C for later published studies, but comparison proved to be diffi­ analysis of gut contents in the laboratory. On some cult due to differences in methodology. Unlike occasions, whole squid were frozen and the alimentary researchers who have studied fish diets (e.g. Berg, tracts were removed in the laboratory, 1979; Hyslop, 1980), most of those who have studied the feeding habits of squids seem to be unaware of the After defrosting of the squid in the laboratory, gut importance of choice of methodology and of how the fullness (stomach and caecum) was estimated by analytical methods may affect the results and conclu­ using a 5-point scale which ranged from 0 (empty) to 4 sions. Consequently, a part of the discussion is (completely full). The gut contents were then trans­ devoted to methodology of feeding habit studies and ferred to a petri-dish and examined under a binocular the pitfalls to be avoided, microscope at magnifications of 6.4-40 x with direct 126 Sci. Council Studies. No.9, 1985 16" 22" hard fragments such as skeletal and jaw elements, parapodia, shell remains and otoliths. Skeletal ele­ 71 0 a 100 km ~ ments of invertebrates are often used for identification purposes and are well described in the literature. On the other hand, descriptions of the form and structure of fish otoliths are relatively scarce. In order to identify 70" fish prey from the otoliths that were recovered from the gut contents of squid, it was necessary to collect oto­ liths from different fish species. Fish were collected with small-meshed bottom and midwater trawls, and 69" the samples were selected to provide as wide a size range as possible for each species. Samples were fro­ zen immediately after capture, and, after defrosting in the laboratory, the standard length (SL) was measured. 68" Sagittal otoliths were removed from the fish and observed under a binocular microscope. Their lengths Fig. 1 Locations where samples of T. sagittatus were obtained were measured and photographs were made to serve along the north coast of Norway. as a ready means of identification. The otoliths were stored in 70% ethanol for use when needed to check illumination. The degree of digestive breakdown of the the identification of otoliths from the gut contents of gut contents was assessed by using a 6-point scale. squid. Furthermore, the linear relationships between with empty guts being classified as 0 and the numbers otolith length and fish length (Table 1) allowed the size 1 to 5 representing degrees of breakdown of the food. of the fish that were consumed by the squid to be The gut contents were generally of a pulp-like or soup­ estimated. like consistency, and the degree of digestive break­ down was most often assessed within the range of 3 to Results 5. This precluded the complete and accurate division of the food remains into specific prey for volumetric or The guts of the squid usually contained very small gravimetric analysis. Consequently, dietary composi­ amounts of well-digested food items, with more than tion was investigated by using indices which were 75% of the guts having a fullness index of 0 to 2. More based on the presence or absence of particular prey than 50% of the guts containing food were assessed as groups (Berg, 1979; Hyslop. 1980). The feeding indices digestive stages 4 and 5. This made identification of are defined as follows: individual prey items difficult and necessitated the use a) "Percentage occurrence" of a given dietary compo­ of small fragments of hard structures for identification nent was calculated as (100 x a)IN, where a is the purposes. The consequence of this is that the contribu­ number of guts containing component a and N is tion of soft-bodied prey to the squid diet is likely to be the total number of guts with food. This index is, underestimated. therefore, a measure of the proportion of squid that has consumed a particular prey. Prey items in the gut contents of squid were classi­ fied into 28 different categories, many of which were b) "Relative frequency of occurrence" of a given prey identified as being representatives of species or genera was calculated by the formula (100x a)La ... z. where of five major animal groups - Chaetognatha, Poly­ a is the number of guts containing component a chaeta, Mollusca, Crustacea and Pisces (Fig. 2). Some and La ... z is the sum of observations of all prey. prey items, such as the chaetognath Sagitta and the This index gives an estimate of how often a particu­ polychaete Eunice, were identified on only one occa­ lar prey appeared in the diet relative to other prey sion, whereas others, such as the polychaete Nereis types. pelagica were found in the guts of some squid from c) For each gut examined, a subjective assessment almost every sample. Two molluscs (Todarodes sagit­ was made of which prey contributed the greatest tatus and Limacina retroversa) were found in the gut biomass to the gut content, and this was termed the contents but the molluscan remains were most fre­ dominant prey. The"dominance" index for a partic­ quently squid suckers and pieces of arms. Among the ular prey was calculated as the proportion of all crustacean prey, the euphausiid Meganyctiphanes dominance values, i.e. (100 x A)IN, where A is the norvegica and the shrimp Pasiphaea sp. were recorded number of guts in which prey a dominated and N is most often, with other crustaceans, such as copepods, the number of guts that were assessed for domi­ being observed sporadically. Fish remains and otoliths nant prey. were found in the gut contents of some squid from each sampling period, and 11 fish species were identi­ As the gut contents of the squid were often well fied. In the samples where pelagic schooling species digested. prey species were usually identified from were identified (Clupea harengus, Mallotus villosus BREIBY and JOBLlNG: Feeding of Todarodes sagittatus off Norway 127 TABLE 1. Linear relationships between otolith length (Ol) and standard length (Sl) and the corresponding coefficients of determination (r') for various fish species. (Sl and Ol in mm.) Size No. Sl = a + b Ol range of Species (mm) fish a b r' Ammodytes tobianus 65-137 42 14.929 50.809 0.93 Clupea harengus 51-255 51 0.240 58.214 0.96 Gadus morhua 26-317 26 5.787 21.993 0.95 Mallotus villosus 62-145 63 25.333 41.780 0.82 Maurolicus mulleri 34-54 32 -7.229 32.091 0.73 Melanogrammus aeglefinus 20-180 20 8.890 17.764 0.98 Micromesisteus poutassou 56-318 27 -25.544 23.170 0.98 Sebastes sp.