Feeding ecology of three species of midwater fishes associated with the continental slope of eastern Tasmania,Australia

The feeding ecology of Maurolicus muelleri, Lampanyctodes hectoris and Diaphus danae was examined from samples collected from continental slope waters of eastern Tasmania between April 1984 and April 1985. A total of 2 232 stomachs was analysed. M. muelleri, L. hectoris and D. danae fed primarily on euphausiids and secondarily on copepods, although larger D. danae (> 60 mm standard length) fed on other lanternfish (chiefly L. hectoris). The diets of M. muelleri and L. hectoris overlapped substantially. Diet overlaps between D. danae and the former species was low, however, due to the large biomass of fish present in D. danae. The prey taxa consumed changed with time of year and predator size. Stomach fullness (feeding intensity) varied seasonally in all three species, but only M. muelleri showed significant diel differences in fullness. The synchronization of the size structure of the predator populations and their feeding intensity, with seasonal variations in preferred prey, is proposed as a mechanism whereby each species maximizes its share of the available food resources.     

Reproductive biology of three species of midwater fishes associated with the continental slope of eastern Tasmania,Australia

The reproductive biology of Lampanyctodes hectoris (Günther, 1876), Maurolicus muelleri (Gmelin, 1789) and Diaphus danae Tåning, 1932, from continental-slope waters of eastern Tasmania, was examined between April 1984 and June 1985. L. hectoris spawned in winter and M. muelleri spawned from late winter to early summer. Apart from one ripe male, no reproductive activity was detected in D. danae; this species may be an expatriate in these waters. Fecundity was positively correlated with standard length in L. hectoris, but not in M. muelleri. The ratio of females to males increased with length in all three species. The spermatozoa of L. hectoris and D. danae are atypical of vertebrates and have no tail.     

Induced anti-predator responses of the green mussel, Perna viridis (L.), on exposure to the predatory gastropod, Thais clavigera Küster, and the swimming crab, Thalamita danae Stimpson

A laboratory experiment was conducted to test whether anti-predator responses could be induced in the green mussel, Perna viridis, on exposure to waterborne cues from two predators, the muricid gastropod, Thais clavigera, and the portunid crab, Thalamita danae, and from damaged conspecifics. There were five treatments in this study. Aquaria of two treatments housed either Thalamita danae or Thais clavigera. Another three treatments housed only intact mussels, broken and intact mussels, or no mussels. No significant differences were found among these five treatments in final shell weight and tissue dry weight. Induced defensive responses were predator-specific. Experimental mussels exposed to waterborne cues of T. clavigera had a higher percentage increase in shell length, height and width, whereas those exposed to waterborne cues of T. danae had a higher percentage increase in shell width and height. Mussels raised in the presence of crabs developed thicker shell at the umbo and lip margin, whereas mussels raised in the presence of gastropods had thicker shell lip. Predator preference for mussels from the control group and for those pre-exposed to waterborne cues of either of the predators was studied. T. clavigera preferentially selected mussels from the control group, whereas no prey preference was found for T. danae, although shell breaking time for mussels exposed to a crab was longer than that for the control. While the present results should be viewed with caution in the absence of a dietary control, this was the first time that prey handling was shown to be affected by defensive changes in the morphology of mussels.Communicated by T. Ikeda, Hakodate     

Heavy metal contamination and physiological variability in the Brazilian mangrove crabs Ucides cordatus and Callinectes danae (Crustacea: Decapoda)

 Physiological studies were made on the crabs Ucides cordatus (L.) and Callinectes danae sampled from populations living in “polluted” mangroves on the southeast littoral of Brazil. Analysis of Cu, Cd, Zn, and Fe of sediments and crab tissues showed interspecific differences in tissue concentrations, and significantly higher levels of Cu, Cd, and Zn in “polluted” populations compared to “unpolluted” crabs living in uncontaminated mangrove in the same geographical area. Individuals of both species from the polluted site showed significantly greater capacities for regulating blood osmotic concentrations at low salinity (9‰). However, U. cordatus showed a reduced hypo-regulatory ability in 34‰S. Differences in ionoregulation were also seen. “Polluted”C. danae showed significantly higher Na/ K-ATPase levels in posterior gills compared to “unpolluted” crabs. Oxygen consumption rates (M˙ _O2) were elevated in U. cordatus, but depressed in C. danae from the “polluted” population. Individuals of both species from this site showed significantly lower O:N ratios, mainly because of an increased net efflux of ammonia. Adenylate energy charge (AEC) values of muscle and hepatopancreas in “unpolluted” and “polluted” populations of both species were not significantly different. These physiological differences are discussed in relation to the known acute physiological and metabolic effects of heavy metals in crustaceans, and interpretated in the light of possible adaptive changes following long-term exposure to contamination. Received: 6 August 1999 / Accepted: 22 June 2000     

Geographic variation in Loligo forbesi in the Northeast Atlantic Ocean: analysis of morphometric data and tests of causal hypotheses

Geographic variation in the squid Loligo forbesi was investigated using multivariate analysis of morphometric and meristic characters in samples of squid taken from 13 localities in the Northeast Atlantic Ocean. Two character sets, body morphometrics and beak morphometrics, indicated similar patterns of variation, with squid from the Azores differing markedly from those on the continental shelf. No consistent pattern was apparent in meristic data. Partial Mantel tests indicated that similarity matrices for morphological data were significantly correlated with distance matrices for (a) geographic proximity, (b) whether the capture site was on the continental shelf or the Azorean bank, and (c) (beak data only) average seasurface temperature at site of capture. Partial Mantel tests on allozyme data for the same individuals support hypothesis (b). The results suggest that L. forbesi in the Azores may reasonably be regarded as a distinct stock, differing significantly from L. forbesi on the continental shelf.     

The trophic link between squid and the emperor penguin <Emphasis Type="Italic">Aptenodytes forsteri</Emphasis> at Pointe Géologie,Antarctica

Cephalopod beaks retrieved from stomachs of dead emperor penguin chicks at Pointe Géologie, Terre Adélie, provide information on taxonomic and size composition of the penguin’s squid diet, on the trophic range of the squid species preyed upon and on the fractional trophic impact of the penguin on the whole food web. Emperor penguins prey upon four squid species (Psychroteuthis glacialis, Kondakovia longimana, Gonatus antarcticus, Alluroteuthis antarcticus) and do not take squid larger than 480 mm mantle length. Larger squid live either below the penguin’s diving range or are beyond its handling capacity. Nitrogen stable isotope ratios indicate that squids cover a range of about two trophic levels (2.5–8‰ δ¹⁵N). The impact of the emperor penguin, however, concentrates on the upper part of this range, about 68% of its squid prey being >6‰ δ¹⁵N. The principal components of the emperor’s diet, fish, krill and squid, differ distinctly in average trophic level. Consequently the trophic position of the emperor penguin changes accordingly with diet composition and may differ by almost one trophic level between different emperor penguin colonies.     

Luminescent flashing in the midwater squids Pterygioteuthis microlampas and P. giardi

The luminescent flashing capabilities and swimming behavior were examined for two species of small midwater squids (Pterygioteuthis microlampas and P. giardi). Recently captured squid were placed in a small aquarium, where their movements and luminescent flashes could be recorded with an image intensifier, a photomultiplier tube and associated equipment. Flashing behavior was initiated by disturbing the squid with an eletrical shock. The following 4 behaviors were observed: (1) a short flash and a rapid departure from the electrode; (2) a short flash and a delayed, slow departure from the electrode; (3) a longer flash persisting after a rapid departure from the electrode; (4) a longer flash when departure was prevented by continuing the shock. Luminescent flashes probably have a general defensive function, discouraging or disrupting an attack by potential predators, and within this context there may exist numerous strategies involving flashes.     

Trophic relations of the cephalopod Martialia hyadesi (Teuthoidea: Ommastrephidae) at the Antarctic Polar Front,Scotia Sea

Samples of the squid Martialia hyadesi were collected aboard two Japanese squid-jigging vessels carrying out commercial fishing trials at the Antarctic Polar Frontal Zone, north Scotia Sea, in February 1989. The dissected stomachs of 61 specimens were classified according to fullness and the contents were examined visually. Identifiable food items included fish sagittal otoliths, crustacean eyes, the lappets on euphausiid first antennule segments and cephalopod sucker rings. The most frequent items in the squid's diet were the myctophid fishes Krefftichthys anderssoni and Electrona carlsbergi, the euphausiid Euphausia superba and a hyperiid amphipod, probably Themisto gaudichaudi. A small proportion of the sample had been feeding cannibalistically. Total lengths of the fish prey were estimated from sagittal otolith size using published relationships. All fish were relatively small; 7 to 35% of squid mantle-length. However, it is possible that some heads of larger fish are discarded by the squid and so are not represented by otoliths in the stomach contents. Over the size range of squid in the sample there was no relationship between size of fish prey and size of squid. Similarly, when the squid sample was divided into groups according to prey categories: crustaceans, crustaceans+fish, fish, cephalopod, there was no evidence that dietary preference was related to squid size. The prevalence of copepod-feeding myctophids in the diet of this squid, which is itself a major prey item of some higher predators in the Scotia Sea, suggests that a previously unrecognised food chain: copepod-myctophid-M. hyadesi-higher predator, may be an important component of the Antarctic oceanic ecosystem.     

Experimental ecology of the temperate scleractinian coral Astrangia danae I. Partition of respiration,photosynthesis and calcification between host and symbionts

Calcification, photosynthesis and respiration of the scleractinian coral Astrangia danae were calculated from the changes in total alkalinity, pH, calculated total CO₂, and oxygen concentration produced by colonies incubated in glass jars. A correction for changes in ammonia, nitrate and nitrite was taken into account and the method evaluated. The fluxes of oxygen and CO₂ were highly correlated (r=0.99). The statistical error of alkalinity determinations was less than 10% of the changes observed in the slowest calcifying samples. Metabolism of polyparium alone was estimated by difference after removal of tissue and reincubation of bare corallum. Zooxanthellae concentration in the polyps was obtained from cell counts made on homogenates of polyp tissue. The calculated photosynthetic rate of the zooxanthellae in vivo was 25 mol O₂ (10⁸ cell)^-1 h^-1 at a light intensity of 120 Ein m^-2 s^-1. In corals having 0.5x10⁹ zooxanthellae/dm² of colony area up to 8% of the total photosynthesis was attributed to the corallum microcosm. Polyp respiration, photosynthesis, and CaCO₃ uptake rates were all much higher than rates previously reported from A. danae, apparently because in these experiments the organisms were better fed. This increased photosynthesis in turn enhanced calcification still further. The symbiosis therefore appears to provide a growth advantage even to fed corals, under the conditions of these experiments.     

Eye development in southern calamary, <Emphasis Type="Italic">Sepioteuthis australis</Emphasis>, embryos and hatchlings

Eye development, optical properties and photomechanical responses were examined in embryos and hatchlings of the southern calamary, Sepioteuthis australis. This species occurs in shallow coastal waters in Australia and New Zealand, and the egg masses were collected in October and December 2004 from Great Oyster Bay, Tasmania. At the earliest developmental stage the eye of the squid was comprised of a hemispherical cup of undifferentiated neural retina, while presumptive iris cell layers and lentigenic precursor cells enclosed a posterior eye chamber. Differentiation of the proximal and distal processes was observed in correspondence with the cornea development and lens crystallization, and occurred before differentiation of the neural retina, which was complete prior to hatching. Longer photoreceptor distal processes were first observed just prior to hatching in the dorsal-posterior retina. After hatching, this difference was much more evident and higher photoreceptor density was found in the central retina. This indicates that the eye of S. australis at this age uses different retina areas for different visual tasks. Optical sensitivity and resolution suggest that juvenile S. australis are diurnal. This study also found functional photomechanical responses of visual screening pigment migration and pupil constriction in S. australis embryos, although complete functionality of the pupil at this stage was uncertain. However, the pupils of squid aged 2 days closed almost completely under bright conditions, showing that photomechanical responses were highly developed in the juvenile squid. These findings indicate that squid embryos are able to perceive visual stimulation, suggesting an early reliance on vision for survival after hatching.     

Feeding ecology and niche segregation in oceanic top predators off eastern Australia

We examined the feeding ecology and niche segregation of the ten most abundant fish species caught by longline operations off eastern Australia between 1992 and 2006. Diets of 3,562 individuals were examined. Hook timer data were collected from a further 328 fish to examine feeding behaviour in relation to depth and time of day. Prey biomass was significantly related to predator species, predator length and year and latitude of capture. Although the fish examined fed on a mix of fish, squid and crustacea, fish dominated the diet of all species except small albacore (Thunnus alalunga) which fed mainly on crustacea and large swordfish (Xiphias gladius) and albacore which fed mainly on squid. Cannibalism was observed in lancetfish (Alepisaurus spp.). Multidimensional scaling identified three species groups based on their diet composition. One group consisted of yellowfin tuna (T. albacares), striped marlin (Tetrapturus audax) and dolphinfish (Coryphaena hippurus); a second group consisted of bigeye tuna (T. obesus), swordfish and albacore; and a third consisted of southern bluefin tuna (T. maccoyii) and blue shark (Prionace glauca). Of note was the separation of mako shark (Isurus oxyrhynchus) and lancetfish from all other predators. Prey length generally increased with increasing predator length although even large predators fed on a wide range of prey lengths including very small prey. Overall, differences in prey type and size, feeding times and depths were noted across the range of species examined to the extent that predators with overlapping prey, either in type or size, fed at different times of the diel period or at different depths. Taken together these data provide evidence for feeding niche segregation across the range of oceanic top predators examined.     

Comparative foraging ecology and ecological niche of a superabundant tropical seabird: the sooty tern Sterna fuscata in the southwest Indian Ocean

Over 6-million pairs of sooty terns Sterna fuscata breed once a year in the southwest Indian Ocean, mostly on three islands of the Mozambique Channel (Europa, Juan de Nova and Glorieuses) and in the Seychelles region. Seasonal reproduction in either winter or summer is the dominant strategy in the area, but non-seasonal reproduction also occurred in some places like at Glorieuses Archipelago. The feeding ecology of the sooty tern was investigated during the breeding seasons to determine whether terns showed significant differences in their trophic ecology between locations. Regurgitations were analyzed to describe the diet of individuals when breeding, and stable isotopes and mercury concentrations were used to temporally integrate over the medium-term of the trophic ecology of both adults and chicks. Overall, the diet was composed of fish, flying squid and fish larvae in different proportions. At Europa and Aride in the Seychelles, where winter reproduction occurs, large epipelagic prey like flying fish or squid dominated the diet. At Juan de Nova, sooty terns reproduce in summer and rely mostly on fish larvae. At Glorieuses (non-seasonal breeding), the diet was intermediate with fish larvae and flying squid being important prey items. The stable-carbon and nitrogen isotope values in blood confirm the differences observed in dietary analysis, and demonstrate different feeding strategies between colonies. δ¹³C values of feathers showed spatial segregation between birds from the Mozambique Channel and the Seychelles region. Terns from the Seychelles had also higher δ¹⁵N values. Feather δ¹³C values also suggest a significant shift from summer to wintering habitat for birds from Juan de Nova. This study emphasizes the high phenotypic plasticity of the species, which may explain its numerical dominance in all tropical waters of the World’s Ocean.     

Genetic evidence of population heterogeneity and cryptic speciation in the ommastrephid squid Martialia hyadesi from the Patagonian Shelf and Antarctic Polar Frontal Zone

Horizontal starch gel electrophoresis was used to investigate levels of genetic differentiation between four samples of the nominate squid species Martialia hyadesi Rochbrune and Mabille, 1889, obtained from regions of the Patagonian Shelf and Antarctic Polar Fron-tal Zone over 1000 km apart. M. hyadesi is an ecologically important South Atlantic ommastrephid squid and it is probable that, in the future, fishing effort will be increasingly directed towards this species. Details regarding the population structure of the species are therefore required. In comparison with the other three samples of M. hyadesi, one of the samples from the Patagonian Shelf (PAT 89II) exhibited fixed allelic differences at 16 of the 39 enzyme loci which were resolved (genetic identity, I=0.51). This high level of genetic differentiation contradicts the apparent morphological similarity between samples, indicating the presence of a cryptic or sibling congeneric species. Deviations from Hardy-Weinberg equilibrium and significant differences in allele distribution were also detected within and between the other three putative M. hyadesi samples, suggesting that the species fails to maintain effective panmixia across its geographical range. The occurrence of both temporal (1986 cf. 1989) and geographic structuring within the species complex is consequently indicated, caused possibly by an overlap of reproductively isolated stocks (stock mixing) outside their respective breeding areas. Low levels of genetic variability were detected throughout the samples examined, estimates of average heterozygosity per locus within the two species detected being in the order of 0.01 and 0.002. These values are discussed in relation to levels of genetic variability reported for other squid species, and in comparison with values typically expected for marine invertebrates.     

Temporal life history plasticity of the Southern Ocean squid <Emphasis Type="Italic">Todarodes filippovae</Emphasis> from waters off Tasmania,Australia

Population dynamics and maturity parameters were analysed for seasonal samples of the oceanic ommastrephid squid Todarodes filippovae from off the coast of Tasmania Australia from 2002 to 2004. Based on assumed daily periodicity in statolith increments, T. filippovae had a life cycle of about a year with the sexually dimorphic females reaching their larger size by predominantly growing faster than males. Due to the small sample size of males, analysis was undertaken on female individuals only. Growth in all samples was best described by a power curve and varied annually, with significantly faster growth in 2001 compared to the subsequent 2 years. Seasonal growth rates also varied with autumn- and winter-hatched squid significantly faster than summer-hatched squid. Spring growth rates were intermediate but not significantly different to the other three seasons. Peak hatching periods occurred in late autumn and early winter. ANOVA revealed a season × year interaction for mantle length and total body weight. Pairwise comparisons showed that the annual differences were likely driven by smaller squid in autumn 2002 compared to autumn samples in the other 2 years. Pairwise comparisons also revealed seasonal differences with winter-caught squid smaller than those from the majority of other samples. There were no seasonal effects on mature female gonad weights but females caught in 2002 had significantly lighter gonad weights than females from the following 2 years. There were no consistent trends among seasons or years in the age structure of mature females. This study revealed the plasticity and flexibility in growth and maturity parameters in this species, with a preference for faster growth during cooler periods.     

Genetic variation in the neritic squid Loligo forbesi (Myopsida: Loliginidae) in the northeast Atlantic Ocean

Horizontal starch gel electrophoresis was employed to investigate levels of genetic differentiation between 13 samples of the neritic squid species Loligo forbesi Steenstrup obtained from throughout the majority of its known geographical range. Six enzyme loci identified in a preliminary study as being polymorphic were screened for variation between samples. No significant differences in allele distribution were detected between any of the samples obtained from the Faroe Bank in the north to Lisbon in the south, suggesting that squid throughout this range in the vicinity of the continental shelf are able to maintain panmixia, and effectively belong to a single population sharing a common gene pool. No clinal variation in allele distribution was detected throughout this range, a result which complements the findings of a detailed morphological companion study of the same individuals. Comparison of this homogenous European continental shelf population with squid from the Azores revealed highly significant (P<0.01) differences in allele distribution at five of the six polymorphic enzyme loci studied. A genetic identity value (I) equivalent to 0.93 over 33 loci was obtained. Analysis of F-statistics suggested migration rates between sites to be as low as one individual per five generations, a rate deemed insufficient under most models to prevent divergence by random genetic drift. The large distance and oceanic depths separating the Azores from continental Europe seem to present an effective barrier to gene flow to L. forbesi, a squid belonging to a family considered to be confined in distribution to relatively shallow, near coastal waters. The two populations of squid in the Azores and along the European continental shelf currently both ascribed to L. forbesi should therefore probably best be regarded as relative subspecies.     

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.     

Contrasting C and N isotope ratios from sperm whale skin and squid between the Gulf of Mexico and Gulf of California: effect of habitat

To investigate feeding variation between populations of an apex oceanic predator, stable isotope ratios of carbon (δ¹³C) and nitrogen (δ¹⁵N) have been compared in skin of female and immature male sperm whales (Physeter macrocephalus) from the northern Gulf of Mexico (GoM) and the Gulf of California (GC). Whale sexes were determined genetically. The δ¹³C and δ¹⁵N values from squid muscle were used from the GC, and from inshore and offshore sites in the GoM. We documented contrastingly lower δ¹³C and δ¹⁵N from whales and squid of the GoM compared with those from the GC. While this difference may be associated with variation in trophic position, geographic variation in biochemical cycling influenced significantly the contrasting isotope values between gulfs. Within the northern GoM, the highly distinct δ¹⁵N values of neritic squid versus mesopelagic squid provide further evidence of habitat specificity in δ¹⁵N.     

Spatial and temporal distributions of copepods to leeward and windward of Oahu, Hawaiian Archipelago

The spatial and temporal distributions of two island-associated copepod species, Undinula vulgaris Dana and Labidocera madurae Scott, were compared to the distributions of two open ocean species, Cosmocalanus darwinii Lubbock and Scolecithrix danae Lubbock, along 28-km windward and leeward transects off the island of Oahu, Hawaii. Samples were taken in September and December 1985 and April and June 1986. A warm, low salinity pool on the leeward side was a prominent feature during all transects except December. The abundances of the two oceanic species did not change significantly between leeward and windward stations, with distance from shore, or between September 1985 and April 1986 samples. As expected, very high abundances of U. vulgaris occurred at some nearshore stations, up to 3 g dry wt m⁻² for adults alone. Calculations of respiratory loss at these densities (0.7 g C m⁻² d⁻¹) suggest a high local productivity would be required to meet these demands. L. madurae, a surface-dwelling species normally restricted to within 1 km of shore, was an effective indicator species of nearshore water movement. It was more common in offshore samples on the leeward transects, rarely being found offshore on the windward side, consistent with prevailing currents and the presence of the leeward warm, low salinity pool. The occurrence of a strong mixing event in April 1986 resulted in L. madurae being distributed throughout the upper 100 m of the water column. The presence of oceanic species close to shore on the windward side also coincided with this wind-driven event. The primary environmental influence on vertical distributions was daytime cloud cover, with U. vulgaris tending to be found shallower on cloudy days. Of the two oceanic species, S. danae exhibited the most pronounced vertical migration, however, vertical distributions were not significantly correlated with environmental factors for either species. The abundant nearshore U. vulgaris population cannot be explained by differences in vertical distribution between it and the two oceanic species that might allow a physical mechanism to concentrate U. vulgaris. A high population growth rate is likely necessary to explain U. vulgaris' dominance. Received: 26 June 1998 / Accepted: 31 March 1999     

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.     

Confirmation of serial spawning in the chokka squid Loligo vulgaris reynaudii off the coast of South Africa

Ovary maturation and spawning in the chokka squid Loligo vulgaris reynaudii were investigated by collecting squid from across their distributional range on the southeastern coast of South Africa between 1992 and 1993. Based on histological examination of the ovarian tissue and the separation of multiple modes in oocyte size-frequency distribution, ovary development was characterised by eight distinct stages. Our study confirms serial spawning in L. v. reynaudii. Received: 9 October 1998 / Accepted: 22 April 1999