Flexible reproductive strategies in tropical and temperate Sepioteuthis squids

A major difficulty confronting the determination of cephalopod reproductive life history is assessing over what portion of the life span an individual is reproductively mature and actively depositing eggs. This paper assesses the potential of the tropical Sepioteuthis lessoniana and two genetic types of the temperate Sepioteuthis australis, to spawn multiple batches of eggs at discrete times throughout the adult life span. This is achieved by histological examination of the ovarian gametogenic cycle and detailed morphological assessments of the reproductive system, in conjunction with other biological information. The genetic type of S. australis found at the northern limits of its Australian distribution showed evidence of a high correlation between body size and quantity of mature eggs, suggesting that eggs may be accumulating to be laid in a single batch. Although maturation was also a size-related process in S. lessoniana and Tasmanian S. australis, oviduct size was not correlated with body weight in mature females, which is indicative of multiple spawning. Further supporting evidence includes relatively low gonadosomatic indices, the heavier weight of the ovary relative to the oviduct, and the feeding activity of mature animals. Mature S. lessoniana and S. australis individuals were present at each location over very wide age and size ranges. In Tasmanian waters, there were distinct seasonal differences in the reproductive biology of S. australis. Summer-caught individuals had much higher gonadosomatic indices and may have been laying larger batches of eggs compared with winter-caught individuals. Summer-caught females also showed a negative correlation between egg size and egg number within the oviduct, suggesting that some individuals were producing fewer, larger eggs and others many smaller eggs. Evidence suggests that considerable flexibility is inherent in the reproductive strategy of both S. lessoniana and S. australis. Received: 11 May 2000 / Accepted: 12 September 2000     

Vertical distribution and feeding patterns of midwater fish in the central equatorial Atlantic II. Sternoptychidae

The vertical distribution of seven sternoptychid species was examined from RMT 1+8 samples collected aboard R. V. Meteor in March-April 1979 and from Royal Research Ship R.R.S. Discovery in July 1974 in the central equatorial Atlantic. During daytime sternoptychids occupied depths between 200 and 1250 m, with Sternoptyx pseudobscura living deepest, centering between 800 and 900 m, and Argyropelecus sladeni most shallow, aggregating predominantly at 300 and 400 m. They are all considered limited or partial migrants, ascending only some 100 and 200 m towards the surface at night. Only A. sladeni was observed to enter the epipelagic zone (0 and 200 m).-Feeding patterns were investigated from stomach content analyses of Sternoptyx diaphana, S. pseudobscura, Argyropelecus sladeni and A. affinis. Additional stomach contents were analysed from samples of S. diaphana, A. hemigymnus and A. olfersi collected in June 1985 from F.R.V. Walther Herwig in the temperate NE Atlantic at 46°N, 17°W by means of the Engel Trawl. The food spectrum of the six species is generally described, and additional dietary evidence regarding calanoid copepod prey is provided for four of these taxa. All sternoptychid species investigated were planktivorous, feeding predominantly on copepods and ostracods, except for the largest size class, which preyed heavily on euphausiids and amphipods. The relationship of predator size towards prey type and prey size is analysed for both Sternoptyx species. Of these, S. pseudobscura in particular exhibits taxonomic selectivity towards polychaete prey. The diet of both species of Sternoptyx included a number of epipelagic or even neustonic calanoid copepod species which contributed more than 50% of the total copepod population by numbers. So far it is not known how the predators find access to prey organisms of the upper 200 m, as netfeeding is considered unlikely. Cyclopoid copepods of the genus Sapphirina were observed as dietary component particular of S. diaphana.     

Vertical distribution and feeding patterns of midwater fish in the central equatorial Atlantic

Fishes and zooplankton were obtained (March–April 1979 and partly in August 1974) from 45 hauls taken during the day and at night in the central equatorial Atlantic between Latitude 3°N and 2°S from the surface to 1250-m depth, using the RMT 1+8, a combined opening-closing plankton and micronekton trawl. The vertical distribution of 30 myctophid species is described. All species migrate in a diel pattern, Ceratoscopelus warmingii and Lampanyctus photonotus down to at least 1250 m. During daytime most species aggregated at 400-to 700-m depth, therefore only partly occupying the depth of the Deep Scattering Layer (400 to 500 m at 15 kHz). The feeding patterns of seven of the most abundant species were compared, with a total of 1 905 stomach contents being analysed. All seven species are regarded as opportunistic predators, which feed predominantly during the night on calanoid copepods. A total of 66 species of calanoid copepods were identified among the prey items, with smaller species definitely being in the minority. Stomachs of C. warmingii (700 to 1 250 m depth) and Lepidophanes guentheri (500 to 900 m depth) from daytime samples contained copepod species restricted to the upper 150 m of the water column, including Undinula vulgaris, Nannocalanus minor, and Euchaeta marina, thereby confirming an extended vertical migration of predators. Differences in diet and preferences between species in their total food spectrum are described.     

Sinking rates and dissolution of midwater fish fecal matter

The sinking rates of fecal matter from 7 southern California midwater fish species were investigated. Feces were obtained from 162 specimens of Stenobrachius leucopsarus, Triphoturus mexicanus, Leuroglossus stilbius, Lampanyctus ritteri, Argyropelecus affinis and Parvilux ingens, which were collected in the Santa Barbara and San Clemente Basins between 1977 and 1979. In addition, feces obtained from 6 laboratory-maintained specimens of the midwater zoarcid Melanostigma pammelas were used for repeated sinking-rate measurements. The mean of the measured sinking rates for all species was 1.19 cm s^-1 (1 028 m d^-1), which is much higher than the known descent rates of euphausiid and copepod fecal pellets and of most other particulate organic detritus. Dissolution characteristics were also investigated for fecal matter from 4 species collected by the same series of net hauls: S. leucopsarus, T. mexicanus, A. affinis, and Sternoptyx obscura. The release of dissolved organic compounds from this material is low and does not represent a significant output during the relatively short time required to sink through the water column. These findings suggest that midwater fish fecal matter may represent a major source of organic transfer between the pelagic community and the benthos.     

Dietary study of the midwater polychaete Poeobius meseres in Monterey Bay,California

This study presents the first quantification of the diet of a gelatinous midwater organism on a temporal basis. Using the Monterey Bay Aquarium Research Institute's remotely operated vehicle Ventana, regular collections of the polychaete Poeobius meseres (Heath, 1930) over a 1 yr period (October 1990 to November 1991) in Monterey Bay yielded intact organisms for the study of feeding behavior and quantitative analysis of stomach contents. In situ observations showed P. meseres feeding in two different ways: (1) by deployment of a mucus web in the water column that passively collects particles for consumption; and/or (2) by grasping detrital material in the water column with its ciliated tentacles. Stomach-content analyses showed that P. meseres is primarily coprophagic, its diet being dominated by fecal pellets from euphausiids and copepods. These fecal pellets appear to provide P. meseres with essentially all its carbon. Although fecal pellets were the most important food item volumetrically, P. meseres also consumed large numbers of diatoms and small numbers of dinoflagellates, chrysophytes, radiolarians, foraminiferans and eggs. The diet of P. meseres appears to reflect primary productivity in the surface waters, with different food items predominant in the diet at different times of the year. Pennate diatoms were most abundant in the diet during the fall, centric diatoms were most abundant during the sumnier, and fecal pellets during the winter. The composition of P. meseres diet suggests that this and other midwater gelatinous organisms have a significant role in the remineralization of particles as they sink from the surface to the deep sea.     

Oxygen consumption of midwater fishes and crustaceans from the eastern Gulf of Mexico

Oxygen consumption was measured as a function of temperature, oxygen partial-pressure (P_O2)and species depth of occurrence for twenty-three species of midwater fishes and crustaceans collected from the eastern Gulf of Mexico from June 1981 to July 1985. Q₁₀s (7° to 20°C) of 3.90 and 3.24 were recorded for myctophid and non-myctophid fish groups, respectively, while values of 2.22, 2.19, 2.19 and 2.54 were calculated for sergestid, penaeid, carid and euphausiid crustacean groups, respectively. Q₁₀s were consistent for species within each group. All of the species tested regulated their oxygen consumption to P_O2levels normally encountered within the eastern Gulf. Values of critical partial pressure (P_c) ranged from 20 to 40mm Hg and increased slightly with increasing temperature and respiration rate. Declining respiration with increasing minimum depth of occurrence was primarily a function of temperature alone. Changes in size, dry weight and water content contributed only a small fraction to the observed decrease. This finding contrasts with studies from the eastern Pacific Ocean, where temperature is a minor contributor to changes in respiration rate with depth.     

The culture and maintenance of the loliginid squids Sepioteuthis sepioidea and Doryteuthis plei

A technique for rearing the loliginid squids Sepioteuthis sepioidea and Doryteuthis plei is reported. Specimens of the former were reared from eggs to sexually mature adults, and maintained for a maximum of 146 days; adult D. plei were maintained for 38 days. Choice and quantity of food was most important for the survival of all sizes, particularly young squid. Newly hatched specimens thrived on Mysidium columbiae. Both species fed at a rate of 30 to 60% of their body weight daily; starvation occurred when intake fell below 10 to 15%. Food conversion efficiency averaged between 10 to 20%. Growth was rapid and steady. S. sepioidea grew to a maximum of 105 mm and 77 g in less than 5 months; D. plei grew an estimated 20 mm/month. Experimental data indicate a lethal minimum salinity for both species at about 27. Lethal minimum and maximum temperatures for young S. sepioidea are 17.5° to 18.0°C, and 32.5° to 33.0°C, respectively. Young consume 0.64l O₂/mg wet weight/h. Opaque tanks, with a semi-natural bottom substrate and special ultra-violet (UV) illumination, are advantageous for rearing and maintenance.Contribution No. 1305 from the Rosenstiel School of Marine and Atmospheric Science, University of Miami, USA. This work was supported in part by the National Science Foundation Grant GB-5729 X. This paper constitutes part of a doctoral dissertation on squid rearing, maintenance and behavior. Aspects of the behavioral studies will be published elsewhere.     

Triacylglycerols and wax esters in the lipids of deep midwater teleost fishes of the Southern California Bight

Total lipid contents and the proportions of triacylglycerols and wax esters were examined in 23 species of deep-sea fishes collected between 1977 and 1980 in the Santa Catalina, San Clemente and San Pedro Basins off Southern California, USA. Mean total lipid content ranged from 1 to 22% of wet weight. Triacylglycerols accounted for 1 to 91% of total lipids present and were more abundant than wax esters in the lipids of all but a single species. Wax ester contents ranged from 0 to 24% of total lipids but in 20 species were 10% of the lipids. These results do not substantiate the previously suggested idea that wax esters are important constituents of the lipids of deep-sea fishes. Moreover, the hypothesis that wax ester content increases with increasing depth of occurrence is not supported. To assess the importance of the lipids present in bouyancy regulation, the water content and swimbladder inflation of these species were also examined. Neither total lipids nor wax esters appear to play an important role in buoyancy regulation in the majority of these fishes. Eight species have gas-filled swimbladders. Of the species lacking inflated swimbladders, 7 have watery bodies (87 to 95% H₂O) and 6 with low lipid (8.3%) and water (85%) contents may need to swim continuously to maintain their position in the water column. Lipids, largely in the form of triacylglycerols, may be important in the buoyancy of three species with lipid contents>10% of wet weight. Two of these species also possess inflated swimbladders. The reasons for the production of large quantitites of was esters by certain deep-sea teleosts, especially gonostomatids and myctophids, remain unknown.     

Chemical composition,metabolic rates and feeding behavior of the midwater ctenophore Bathocyroe fosteri

Individuals of the midwater ctenophore Bathocyroe fosteri (0.01 to 1.6 g dry weight, DW) were collected from Bahamian waters by the submersible Johnson-Sea-Link during May and September/October 1983 and October/November 1984 from 530 to 700 m depth. Metabolic rates were measured and showed oxygen consumption to be in the range of 0.01 to 0.18 mg O₂ g^-1 DW h^-1 at temperatures ranging from 9° to 12°C. Ammonium excretion (0.01 to 0.14 g-at N g^-1 DW h^-1) was typically low. Energy expenditures estimated from respiration data (ca. 7% body C d^-1) indicated that one to three midwater crustacean prey (ca. 150 g C d^-1) could provide the daily maintenance ration required by a 40 mm ctenophore. These metabolic characteristics complemented in situ observations of poor locomotor ability and passive feeding behavior.     

Distribution and community structure of midwater fishes in relation to the subtropical convergence off eastern Tasmania,Australia

Midwater fishes were sampled at night to a depth of 400 m in oceanic waters east of Tasmania, Australia, in May/June of 1992, 1993 and 1994. We examined whether there were differences in the distribution and density of these fishes in relation to the subtropical convergence (STC) separating East Australia Current (EAC) water from subantarctic water (SAW) to the south. A total of 23999 fishes from 107 taxa and 43 families were identified. Four new records were identified from the study area. Myctophids and stomiatoids were the main fish taxa captured. We found no evidence of increased density of the combined fish catch in any particular area. However, differences between areas in the density of individual species were noted, although no species was confined to any one area. Ceratoscopelus warmingi, Lobianchia dofleini and Vinciguerria spp. were most abundant in the EAC, whereas Lampichthys procerus was most abundant in the SAW. Only Diaphus danae was found in significantly higher numbers in the STC. Multivariate analysis revealed that community structure in the EAC was significantly different from that of the SAW but not from the STC. Further, when separated by depth, the shallow EAC group was significantly different in all group comparisons bar that with the shallow STC. Our data indicated that the thin EAC layer above 200 m could be distinguished by its fauna from the SAW. However, we could not detect a separate community in the STC. We conclude, therefore, that the STC is not an area of increased micronekton abundance, nor does it contain a distinct community, during the Australian autumn/winter.     

Eyes and extraocular photoreceptors in midwater cephalopods and fishes: Their roles in detecting downwelling light for counterillumination

The means of detecting downwelling light for counterillumination in several midwater animals has been examined. Eyes and extraocular photoreceptors (drosal photosensitive vesicles in the enoploteuthid squid Abraliopsis sp. B and pineal organs in the myctophid fish Myctophum spinosum) were alternately exposed to overhead light or covered by a small opaque shield above the animal and the bioluminescent response of the animal was monitored. Covering either the eyes or the extraocular photoreceptors resulted in a reduction in the intensity of counterillumination. Preliminary experiments examining the bioluminescent feedback mechanism for monitoring intensity of bioluminescence during counterillumination in the midwater squid Abralia trigonura indicated that the ventral photosensitive vesicles are responsible for bioluminescent feedback.     

Behaviour of the institute of oceanographic science's rectangular midwater trawls: Theoretical aspects and experimental observations

The behaviour of two rectangular midwater trawls, the RMT 1+8 and the RMT 1+8M, is described. The RMT 1+8 consists of a pair of rectangular nets contained within the same frame-a small mouth area RMT 1 of mesh size 320 m and a larger RMT 8 of mesh size 4.5 mm. The RMT 1+8M is a multiple version of the RMT 1+8, combining three RMT 1s and three RMT 8s within the same frame. An expression is developed to describe the dependence of mouth angle on speed for the RMT 1+8 when towed horizontally. Theoretical estimates of the changes in mouth angle with speed agree well with experimental observations. Within normal towing speeds the RMT 1+8 is sensitive to changes in speed; this sensitivity is especially marked in the RMT 1. Between speeds of about 2.6 and 8.0 knots the RMT 8 does not alter its mouth angle with changes in speed; this behaviour is not shown by the RMT 1—at least within a practical speed range. The performance of the RMT 1+8M cannot be completely described by the theory developed for the RMT 1+8 and possible explanations for this are discussed. The multiple net is considerably less sensitive to changes in towing speed; all three nets of each type have similar mouth angles at a constant speed. Variations in warp load with speed for both trawls are described.     

Food availability as a selective factor on the chemical compositions of midwater fishes in the eastern North Pacific

In September and October 1980 we examined the relationships between food availability, depth, and chemical composition among 12 midwater fish species, from three adjacent areas of the eastern North Pacific: the eastern gyre, the California Current, and the transitional region between them. By comparing trends in chemical composition across a geographical productivity gradient, the influence of food availability could be examined both with and without depth as a dependent variable. In general, caloric density, lipids, and water content showed consistent trends along both vertical and horizontal gradients of food availability. Lipids and caloric contents were lowest among bathypelagic species and among fishes from the gyre. Water content was highest in the gyre and among deeper-living species. While protein content as a percentage of the total wet weight declined with depth, there was no clear trend among mesopelagic species between stations. Protein as percent of the total organic content showed no decrease with depth, but was significantly higher in the gyre among mesopelagic species. These findings suggest that compositional trends in water, lipids, and caloric content are correlated with food availability, regardless of any depth-related factors. In contrast, protein did not vary consistently with food availability and thus may be linked to other factors.     

发光细菌生物活性的调控方法

为维持发光细菌发光强度的稳定性,推进发光细菌毒性测试技术应用于在线监测和分析,以明亮发光杆菌和鳆鱼发光杆菌为对象,通过添加各种保护剂,将培养至对数生长期的菌液离心,重新悬浮于脱脂牛奶溶液冷藏(5℃),比较脱脂牛奶菌悬液冷藏、冻干粉复苏后即时冷藏以及新鲜菌液直接冷藏3种方法的调控效果.结果表明,脱脂牛奶菌悬液冷藏7 d后复苏,相对发光率达到93%.该方法明显提高了发光细菌生物活性的稳定性,对于提高在线毒性监测仪连续运行时间有参考价值.     

Phylogenetic diversity of epibiotic bacteria in the accessory nidamental glands of squids (Cephalopoda: Loliginidae and Idiosepiidae)

Bacterial communities were identified from the accessory nidamental glands (ANGs) of European and Western Pacific squids of the families Loliginidae and Idiosepiidae, as also in the egg capsules, embryo and yolk of two loliginid squid species, and in the entire egg of one idiosepiid squid species. The results of phylogenetic analyses of 16S RNA gene (rDNA) confirmed that several phylotypes of α-proteobacteria, γ-proteobacteria and Cytophaga–Flavobacteria–Bacteroides phylum were present as potential symbiotic associations within the ANGs. Several identified clones were related to reference strains, while others had no known close relatives. Gram positive strains were rare in loliginid squids. Several bacterial groups may play important roles in the function of the ANGs, such as production of the toxic compounds involved in egg protection and carotenoid pigments. Within the eggs, no bacteria were associated with embryo or yolk of Loligo vulgaris and Sepioteuthis lessoniana, but α- and γ-proteobacteria were present in the egg capsules. Most bacterial strains detected in the egg capsules were the same as those found in the ANGs. The cephalopods of temperate regions (European cuttlefishes and the squid L. vulgaris) appear to be associated with one Agrobacterium strain (Agro2) while tropical-subtropical strains (Asian and Australian loliginids) are associated with Silicibacter-related strains, suggesting a biogeographic clustering for the Agrobacterium-like strains.     

Bioluminescence in the Monterey Submarine Canyon: image analysis of video recordings from a midwater submersible

Video images of bioluminescence were recorded in situ during a 1985 study of the midwater environment of the Monterey Canyon, using a single-person, untethered submersible. Gelatinous organisms were responsible for the most brilliant bioluminescent displays, often exhibiting elaborate kinetics in response to mechanical stimulation. Images of bioluminescent displays recorded from identified organisms are shown and display patterns are described. All bioluminescence emission spectra from captured specimens were blue, with peak emissions between 460 and 494 nm. Image-analysis of recordings of mechanically stimulated bioluminescence revealed source densities between 43 and 175 m^-3 and intensities between 2.5 and 37.3 W sr^-1 m^-3. The predominant display type at all depths studied (between 100 and 560 m) was luminous secretions. Despite high intensities of mechanically stimulated bioluminescence, no spontaneous light production was recorded in the absence of mechanical stimulation.     

Metabolic characteristics of midwater zooplankton: Ammonia excretion,O:N rations,and the effect of starvation

The nature of protein catabolism in a wide range of species of midwater zooplankton was investigated. The weight-specific ammonia excretion rates (g NH₃–N g^–1 dry wt h^–1, y) decline exponentially with minimum depth of occurreece (MDO, x), y=163.4 x^{–0.479±0.212} (95%ci) (CI=confidence interval), when temperature is held constant. The change in ammonia excretion can be partially explained by the decrease in percent protein (%P) with MDO, %P=80.17 MDO^{–0.148±0.122 (95%ci)} The atomic O:N ratio of freshly caught zooplankters ranged from 9.1 to 91, with most measurements between 9 and 25. Detailed studies were carried out on the response of one of the species studied (Gnathophausia ingens) to starvation (28 d). After 14 d of starvation the average ammonia excretion rate declined by more than 75% to less than 1 g NH₃–N g^–1 wet wt h^–1, although the average oxygen consumption declined by only 13% within the first 7 d of starvation and then remained stable. This differential response of oxygen consumption and ammonia excretion to starvation resulted in an increase in the average O:N ratio of starved animals from an initial 33 to 165 after 21 d. The average O:N ratios of fed mysids remained below 38 during the experiment. G. ingens maintains a relatively uniform metabolic rate during starvation by relying more heavily on its large lipid stores than when being fed.     

A comparative study of metabolic energy expenditure in the limpets Patella cochlear,P. oculus and P. granularis

Oxygen consumption of 3 species of Patella was measured in air and water at various temperatures. Measurements at constant temperature over a full tidal cycle showed no tidal or light-dark rhythms. Measurements under conditions simulating natural tidal, temperature and day-night cycles allowed calculation of daily respiratory energy budgets. P. cochlear occurs low on the shore, but experiences a food shortage due to intense intraspecific competition. Its rate of respiration is moderate, but metabolic expenditure is kept low because exposure to air is brief and body temperatures seldom rise above 23°C. P. cochlear has a respiratory rate-temperature (R-T) curve which peaks at 20°C and forms a plateau between 20° and 32.5°C. The midshore P. oculus has abundant food and adopts an exploitative strategy. Growth rate is very high, and this high turnover of energy is linked with a high metabolic rate, high Q10 (temperature coefficient) values, high body temperatures during the day-time low tide, and a respiratory R-T curve peaking at 32.5°C. Small P. oculus occur mainly in intertidal pools and respire faster in water, while larger individuals occur on bare rocks and respire faster in air over the upper temperature range. In contrast, the upper-shore P. granularis has little food, and conservation of energy is essential, particularly as its growth rate is moderate and its reprocurve output high. Respiratory losses are reduced by suppression of the R-T curve and low Q10 values, resulting in relative independence of temperature. Small P. granularis occur low on the shore and respire slower in water. Larger individuals occur at high levels due to migration, and respire slower in air. This further reduces respiratory energy losses. The patterns of respiration in these 3 species are thus related to food availability, resulting in exploitative or conservationist strategies.     

Growth rhythms in the shrimp Penaeus vannamei and P. schmitti

Growth increments of the penaeid shrimp Penaeus vannamel (Boone) and P. schmitti (Burkenroad) grown in commercial production ponds in Ecuador and Colombia were measured over a 4 mo period between October 1987 and January 1988. P. vannamel grown from wild caught seed in Ecuadorian ponds exhibited a growth rhythm which significantly coincided with lunar phases. Growth increments of at least 1 gwk^-1 occurred around a new and full moon, while growth increments of less than 1 gwk^-1 occurred during first and last quarters. P. vannamel transported from Ecuador as nauplii, raised in Colombian hatcheries and on grown in Colombian ponds exhibited a cyclic pattern in wet weight increments but not correlation to lunar cycles was observed. P. schmitti showed no cyclic pattern in growth. The two species under consideration are indigenous to two different, and quite unique areas displaying different environmental characteristics. The link between the growth pattern, biological and lunar cycles is discussed.     

Persistent locomotor rhythmicity in the prawns Penaeus indicus and P. monodon

The locomotory activity of adult Penaeus indicus Milne Edwards and P. monodon (Fabricius) collected from the Vellar estuary, South India, in 1983 was monitored continuously for a period of 40 d in the laboratory. Both species exhibited persistent circatidal and circadian rhythmicity, the former displayed peak activity during predicated times of high water in their original habitat. The latter exhibited peak activity during periods corresponding to in situ night-time, with almost no activity during the day, except at the time of high tide. The tidal and daily rhythms were evident for only 3 to 4 d and 5 to 6 d, respectively, under laboratory conditions. The amplitude of the rhythms displayed variations in each semilunar period, reflecting the synchrony and dissynchrony of the tidal and daily rhythms. Activity was correlated with phases of the moon, peaking at times of new and full moon, and showing a scattered pattern with smaller peaks during the first and third quarters.