ALBERT

Description: Males of Eledone cirrhosa grow to a size little over 600 g and normally have well-developed, and presumably active, reproductive organs from about 200 g upwards. Total weight of the genital bag is well correlated with total body weight (r= 0·906). Growth of the testis precedes that of the spermatophoric sac, and the size of neither of these reproductive components is predictable from body weight. The sizes of these organs and the estimated number and length of stored spermatophores are given for 100 g intervals of total body weight. No evidence was obtained for a seasonal trend in male maturity.

Description: We present chemical data on magmatically heterogeneous pyroclastic deposits of late Quaternary age erupted from zoned magma systems underlying Tenerife (Canary Islands), Sao Miguel and Faial (Azores), and Vesuvius. The most fractionated magmas present at each centre are respectively Na-rich phonolite, trachyte, and K-rich phonolite. Within any one deposit, chemical variation is either accompanied by changes in the phenocryst assemblage (petrographic zonation) or is largely manifested in trace element abundances, unaccompanied by any petrographic change (occult zonation). Zoning is analogous to that in calc-alkaline systems where the most fractionated products are high-silica rhyolites. When a range of magma types are considered, a correlation emerges between roofward depletion of trace elements (especially REE) in the zoned system and compatability of those same trace elements in the accessory phenocryst phases present. Thus, allanite- or chevkinite-bearing rhyolitic systems are light-REE depleted roofwards, the sphene-bearing Tenerife system is middle-REE depleted roofwards, the melanite-bearing Vesuvius system is heavy-REE depleted roofwards, while the Azores systems, which lack these phases, display roofward REE enrichment. Therefore, the behaviour of trace elements may in each case be explained by fractionation of observed phenocryst assemblages. The resemblance between features of zoned magma systems and published work on the dynamic consequences of cooling saturated aqueous solutions prompts us to suggest that sidewall crystallization and consequent boundary-layer uprise to form a capping layer at top of the system may be a plausible mechanism for the generation of both petrographic and occult zonation. Reverse zoning occurs among the first-erupted tephra of some deposits, demonstrating that the most highly differentiated magma available is not always the first to be tapped during an eruption from a zoned system.

Description: Changes in the relative size of the ovary, oviducal glands and eggs are described for Eledone cirrhosa captured from the North Sea off Aberdeen over a 3 year period (N = 488). The analysis is based only on freshly caught animals, excluding those held in aquarium conditions (〉 5 days). Ovary enlargement and egg size estimates are used as indices of sexual maturity. Between 0–15% and 18–95% of total body weight is contributed by the ovary. Maximum egg length in the ovary ranges up to 7 mm. On these criteria, sexual maturation typically occurs at body sizes between 400–1000 g although some animals of 1000–1200 g are found showing no evidence of ovary enlargement. The majority of the monthly sample is always immature but maturation can apparently occur at almost any time of the year. Increase in mean ovary index and mean values for egg size are strongly seasonal and indicate a peak incidence of sexual maturity over 2–3 months in the July-September period. Spawning is presumed to follow within 1 month. Estimates of the fecundity of the females, based on the egg sample from the ovary, range from 2·2 × 103 to 55 × 10 3 eggs with a mean of 11 × 10 3 and a mode of 7·5 × 10 3 eggs.

Description: Female Eledone cirrhosa held in aquarium conditions for periods of time of five daysand over show relatively enlarged ovary sizes. Values for ovary index considerably exceed thoseof freshly caught animals and the incidence of the final stage of maturity, in which eggs pack the oviducts, is greater. A comparison of maturity indices for fresh and aquarium males was inconclusive. The range of factors associated with aquarium conditions is briefly reviewed and it is concluded that studies of cephalopod reproductive maturation must distinguish fresh and aquarium animals. Introduction External factors effective in inducing sexual maturation in cephalopods have been suggested several times. The influence of the absence of light has been implicated since the experiments of Wells & Wells (1959) showed that blinded Octopus vulgar is matured precociously. An effect of short day length in stimulating the optic glands of Sepia has been found by Defretin & Richard (1967) and Richard (1967) but this is not clearly the case for Octopus (Buckley, 1977). Octopuses kept in aquarium conditions for lengthy periods are said to have larger relative gonad sizes than those fresh from the sea (Wells & Wells, 1975). One of the factors associated with aquarium conditions is often a degree of starvation, and this circumstance alone is held to be a factor in inducing precocious sexual maturation in Eledone (Mangold & Boucher-Rodoni, 1973). In the course of recent studies on the growth and reproduction of Eledone cirrhosa from the North Sea (Boyle & Knobloch, 1982,1983,1984) animals which had remained in aquarium conditions for 5 days or over were separated from the analysis.

Description: A new species of a monogenean Isancistrum subulatae (Gyrodactylidae) has been discovered on the arms and tentacles of the cephalopod mollusc Alloteuthis subulata at Plymouth and I. loliginis, on the gills of the same host, has been re-discovered for the first time since its original description in 1912. I. subulatae, like other gyrodactylids, is viviparous, and has been shown by experiments to transfer to new hosts by contagion. In nature such transfers probably take place during copulation of the hosts and since the parasite may occur in numbers of several thousands per host, it may thereby constitute a venereal disease.

Description: The growth of the octopus Eledone cirrhosa has been studied in a population from the North Sea off Aberdeen. Data are presented for the growth of individuals isolated in aquarium conditions; the growth of size classes in thefieldpopulation; and preliminary information on the growth relationships of gonad, somatic, cardiac and brain components of the body. At 15 °C Eledone cirrhosa is capable of growing from 10 to 1000 g in 270 days. From octopuses which feed readily in captivity, weight specific growth rates of up to about 3–5 % day-1 for animals of 100 g body weight are recorded, falling to a maximum of about 1–5 % day-1 at body sizes above 500 g. Females stop growing when sexually mature, but in the sample captured they were consistently larger than males, a feature which may account for the 7:1 bias towards the incidence of females. On a wet-weight basis, the mean food incorporation into growth is 37 % of the food ingested, which is 49% of the gross weight of crabs killed. Field data for 1978/79 suggest that animals recruited to the population at the beginning of the year grew steadily until December, overwintered without growing, then grew rapidly for several months in the subsequent year before disappearing from the samples. The estimated average age of those animals and by implication, the life span, is 20 months.

Description: The ommastrephid squids are large active animals occurring in most of the world's oceans. Luminous organs or bioluminescence have been observed only in members of the subfamily Sthenoteuthinae, containing the genera Ornithoteuthis, Symplectoteuthis (= Eucleoteuthis), Hyaloteuthis, Ommastrephes and Dosidicus. The light organs of Ommastrephes pteropus are small sub-spherical bodies randomly distributed over the ventral surface of the mantle, head, arms and tentacles (Roper, 1963) and are aggregated dorsally to form a large luminous patch (Clarke, 1965). Relatively little is known about the organs, capabilities and biochemistry of luminescence in cephalopods (Harvey, 1952; Herring, in Press), and the size of the light organ and availability of O. pteropus provide an unusual opportunity for such studies. Although among the molluscs the luminescent systems of the gastropod Latia and the bivalve Pholas have been partially characterized (Shimomura & Johnson, 1968; Henry, Isambert & Michelson, 1970, 1973) the only cephalopod system which has been investigated to date is that of the enoploteuthid Watasenia scintillans (Goto et al., 1974; Inoue et al., 1975). This investigation examines the anatomy and biochemistry of the dorsal light organ of O. pteropus, which differs markedly in these respects from the brachial organs of Watasenia.

Description: Squids (teuthoids) fall into two distinct groups according to their density in sea water. Squids of one group are considerably denser than sea water and must swim to stop sinking; squids in the other group are nearly neutrally buoyant. Analyses show that in almost all the neutrally buoyant squids large amounts of ammonium are present. This ammonium is not uniformly distributed throughout the body but is mostly confined to special tissues where its concentration can approach half molar. The locations of such tissues differ according to the species and developmental stage of the squid. It is clear that the ammonium-rich solution are almost isosmotic with sea water but of lower density and they are present in sufficient volume to provide the main buoyancy mechanism of these squids. A variety of evidence is given which suggests that squids in no less than 12 of the 26 families achieve near-neutral buoyancy in this way and that 14 families contain squids appreciably denser than sea water [at least one family contains both types of squid]. Some of the ammonium-rich squids are extremely abundant in the oceans.

Description: The effect of medium dissolved-oxygen tension on the molar growth yield, respiration and cytochrome content of Beneckea natriegens in chemostat culture (D 0·37 hr-1) was examined. The molar growth yield (Y), the specific rate of oxygen (qo2) and glucose consumption, and the specific rate of carbon dioxide evolution were independent of the dissolved-oxygen tension above a critical value (〈 2 mmHg). However, the potential respiration rate increased with reduction in the dissolved-oxygen tension at values of the dissolved-oxygen tension well above the critical value. Changes in the cytochrome content occurred at dissolved-oxygen tensions well above the critical value. An increase in cytochrome c relative to cytochrome b was observed as the dissolved-oxygen tension was decreased. Reduction of the dissolved-oxygen tension to less than 1 mmHg caused a switch to fermentative metabolism shown by the apparent rise in Y o2 and decrease in the molar growth yield from glucose. At this point the potential respiration rate (q o2) increased to its highest value, while the cytochrome pattern reverted to that observed at dissolved-oxygen tensions above 96 mmHg. There appeared to be no correlation between cytochrome content, potential q o2, in situ q o2, and cyanide sensitivity of the organism at various dissolved-oxygen tensions.

Description: The planktonic foraminifer Globigerinoides sacculifer (Brady) was cultured under two different light intensities and in continuous darkness. High light intensity (HLI = 4oo-soo einsteins/m2/s) resulted in a longer lifespan, a greater number of chambers formed, and a larger final shell size compared with individuals cultured under low light intensity (LLI = 20-50 einsteins/m2/s) or in continuous darkness. Shell growth rates were unaffected by increasing light intensity, but gametogenesis was delayed. Continuous darkness induced a rapid onset of gametogenesis in organisms with shell lengths larger than 250 m. Feeding frequency had a greater effect on growth and reproduction than light intensity under conditions of LLI and HLI, but continuous darkness had an overriding effect on growth and reproduction owing to the rapid onset of gametogenesis which terminated the life of the mother cell. Our previous data indicated that the longevity of G. sacculifer was dependent on feeding frequency, and that G. sacculifer cultured under LLI had a lifespan of approximately 2-4 weeks. Present results suggest that the lifespan can vary from a minimum of 8 days for organisms fed daily in continuous darkness to a maximum of 54 days for organisms fed once every 7 days and maintained in HLI. It is concluded that individual G. sacculifer attain a shell size greater than 6oo ,urn only if they maintain their position in the euphotic zone. Prolonged existence below the euphotic zone would result in premature death or gametogenesis following stunted shell growth.

Description: Terms, dimensions and ratios for statolith description are defined. The form of the calcareousstatoliths in the Teuthoidea, Sepiodea and Octopoda is described by reference to Loligo forbesi, Sepia officinalis and Eledone cirrosa. While statoliths change in form and size during the growth of a cephalopod, the adult form is often characteristic for a species, despite some variation. Description of statoliths is important in studies of the fossil remains of cephalopods lacking calcareous shells, and will probably become important in the taxonomy of living species, in food analysis of cephalopod predators and in the study of deep sea deposits.

Description: Five hundred and twenty-eight specimens of Loligo forbesi Steenstrup, from landings in Horta, Faial, Azores, during the year 1 March 1980 to 28 February 1981, were tudied; 59·3 % were males, 40·7 % females. Of the males 80·2 % were sexually mature, of the females 91·6 %, both sexes showing the highest degree of maturity in spring and he lowest in autumn. The mean dorsal mantle length of the mature males was 56·5 cm, or females 33·5 cm. A weight-length relationship was calculated. he stomachs of 622 specimens were sampled, of which 306 contained food. The prey omponents were studied qualitatively. The main prey was fish (82·0%), of which 0·5 % were horse mackerel, Trachurus picturatus, this being the most important food rganism. Preliminary results of statolith readings are given.

Description: Little work on vertical distribution of cephalopods was possible before the development, in the 1960s, of sophisticated opening-closing devices usable on midwater trawls such as the 10 ft Isaacs Kidd trawl (IKMT; Foxton, 1963; Aron et al. 1964) and the series of rectangular midwater trawls developed by the Institute of Oceanographic Sciences (previously the National Institute of Oceanography) (Clarke, 1969 a; Baker et al. 1973). These developments have resulted in three papers on vertical distribution of cephalopods in the North Atlantic (Clarke, 1969 ft; Gibbs & Roper, 1970; Clarke & Lu, 1974) and one for the Mediterranean (Roper, 1972). The present paper describes the vertical distribution of cephalopods caught at 40° N 20° W, 53° N 20° W and 60° N 20° W in the North Atlantic based upon day and night series of horizontal hauls between the surface and 2000 m using the RMT combination net (Baker et al. 1973).

Description: The present work is part of an analysis of catches made with rectangular midwater trawls (RMTs) in the North Atlantic at about 20°W and at 60°N, 53°N, 40°N (all in Lu & Clarke, 1975), 30°N (Clarke & Lu, 1974), 18°N and 11°N (Lu & Clarke, 1975). The collections were made for the ecological programme of the National Institute of Oceanography, Wormley, England (now part of the Institute of Oceanographic Sciences).

Description: This is one of a series of four papers dealing with vertical distribution of cephalopods in the North Eastern Atlantic at six stations near 20° W and at about 10° intervals from 60°N to 11° N (Clarke & Lu, 1974, 1975 a; Lu & Clarke, 1975). The present study is based upon a series of hauls made at discrete horizons between o and 2000 m with opening-closing nets during both daylight and darkness. The collections were made for the ecological programme of the National Institute of Oceanography, Wormley, Surrey, England (now part of the Institute of Oceanographic Sciences).

Description: Symbiotic luminous bacteria have been described in, and cultured from, a number of species offish and cephalopod. Indeed only in these two groups are extracellular luminous bacteria believed to be utilized as a source of light (see Buchner (1965) and Herring (1978) for references). Despite several earlier investigations of such symbioses in cephalopods the bacteria in these animals have not been adequately identified, nor has the extent of their role been clarified. The ultrastructural relationships between bacteria and the tissues of the squid accessory nidamental gland have been investigated in the non-luminous species Loligo pealei (Lesueur) (Bloodgood, 1977) and Sepia officinalis L. (Van den Branden et al. 1979) but no comparative work on luminous species has been undertaken apart from that on Heteroteuthis dispar (Rüppell), whose photophore does not contain typical luminous bacteria (Dilly & Herring, 1978; cf. Leisman, Cohn & Nealson, 1980). The order Sepioidea contains five families, among which are the two families Sepiolidae and Spirulidae. Though the presence of luminous bacteria is known in some sepiolids (as well as in certain loliginids (order Teuthoidea)) some doubt remains about the source of light in the photophore of Spirula spirula Hoyle. The steady luminescence of this species has prompted speculation that bacteria may be involved (Harvey, 1952). In this paper we compare the anatomy and ultrastructure of the photophores of both Sepiola and Spirula in order to clarify some of these problems.

Description: This is the first detailed analysis of cephalopod beaks from the stomach of a northern bottlenosed whale, Hyperoodon ampullatus (Forster, 1770). The digestive action of many predators barely affects the chitinous beaks of cephalopods and some cetaceans accumulate the beaks in considerable numbers in their stomachs. The present beaks are clean and unbroken. Identification of cephalopod beaks from stomachs of predators such as sperm whales (see Clarke, 1977), seals (Clarke & Trillmich, 1980) and albatrosses (Clarke, Croxall & Prince, 1980) throws considerable light on the biology and relative ecological importance of the species of cephalopods concerned as well as providing useful information on the diet of the predators.

Description: Cephalopod statoliths are paired calcareous stones which lie in cavities, the statocysts, within the skull. They have a form which, though variable, shows promise as a source of criteria for taxonomic and evolutionary studies. As a preliminary to more detailed studies, Clarke (1978) published a description of the form of a generalized teuthoid statolith, coined nomenclature for the various parts and gave a very brief survey of variation of statoliths within the living Cephalopoda. This nomenclature was used in a detailed description of fossilized teuthoid statoliths by Clarke & Fitch (1979). Here, descriptions of the statoliths of the living species Berryteuthis magister (Berry, 1913), Gonatopsis borealis Sasaki, 1923, Gonatopsis (Boreoteuthis) makko Okutani & Nemoto, 1964 and Gonatus fabricii (Lichtenstein, 1818) are given and the fossil Berryteuthis species described in outline by Clarke & Fitch (1979) is compared with B. magister. A statistical analysis of measurements of the statoliths of these five species has been made and the results are presented. This forms the first part of a general description of teuthoid statoliths and similar studies on the Ommastrephidae and the Loliginidae are in preparation.