Reproductive behavior in the squid Sepioteuthis australis from South Australia: ethogram of reproductive body patterns

Squids use a diverse range of body patterns for communication. Each pattern consists of a series of chromatic, postural, and locomotor components that are under neural control and can change within fractions of a second. Here we describe an ethogram of 48 body pattern components from in situ observations of reproductively active Sepioteuthis australis. In addition, we identify the total time and average duration that each component is shown, to a resolution of 1 s. Our results suggest that only a few components (e.g., "Golden epaulettes," "Stitchwork fins," and "Rigid arms") are temporally common, that is, shown for more than 80% of the time. In contrast to the component classification reported for other species of squid, for this species we suggest a classification system consisting of "short acute" (lasting for < 10 s); some of these same components were also classified as "medium acute" (11-60 s) or "chronic" (> 60 s). Several body patterning components were previously unreported, as were some of the combinations observed. The significance of these patterning components is discussed within the context of the associated behaviors of the squid on the spawning grounds.     

Effects of commercial fishing on the population structure of spawning southern calamary (Sepioteuthis australis)

This study explored population dynamics of Sepioteuthis australis on a fine temporal scale before, during, and after a 3-month commercial fishing closure on the summer inshore spawning grounds of Great Oyster Bay, Tasmania, Australia. An abrupt change in male size (mantle length) and population sex ratio after the re-opening of the commercial fishery suggests that fishing alters the population structure on the spawning beds from a ‘natural’ structure which is highly biased towards males, to a more even ratio of males to females. Although jigs are taking a representative sample of the squid population using the spawning beds at any one time, the fishery is apparently still effectively selective for males, potentially as a function of differential spawning movements of the two sexes. Increased fishing pressure over the past 5 years had been correlated with a change towards a highly male-biased sex ratio on the spawning beds; however the current study suggests that increased fishing pressure in recent years may not have reduced the proportion of females. Instead, the inter-annual change in sex ratio may reflect changes in the degree of protection from fishing, with progressively longer closures allowing time for more males to accumulate on the beds and therefore, for the proportion of males to increase (as opposed to a decrease of females). As fishing is selective for males, fishing throughout the spawning season could potentially modify the process of sexual selection and the mating behaviors of the individuals within the spawning population, highlighting the need for closures over this crucial period. Additionally, examinations and comparisons of squid population structure need to be interpreted in light of fishing pressure and broader movement patterns.     

The effects of egg position, egg mass size, substrate and biofouling on embryo mortality in the squid Sepioteuthis australis

Using a combination of laboratory and field investigations, this study examined embryo mortality in the southern calamary Sepioteuthis australis as a function of egg mass size, the substrate upon which the mass is attached, the position of the embryo within the mass, and the degree of biofouling. Egg mass size ranged from 2 to 1,241 egg strands, however most masses consisted of 200–299 strands. Small egg masses (<300 strands) were generally attached to soft-sediment vegetation (Amphibolis antarctica, Heterozostera tasmanica, Caulerpa sp.), whereas larger masses (>300 strands) were either securely attached to robust macroalgae holdfasts (Ecklonia sp., Marcocystis pyrifera, Sargassum sp.) or unattached. Rates of embryo mortality were highly variable ranging from 2 to 25%. Both laboratory and field results indicated a positive relationship between egg mass size and embryo mortality. Larger, unattached egg masses contained twice as many dead embryos than those securely attached to a substrate. Mortality rates were significantly affected by the embryos’ relative position within the mass and were highest in embryos located near the attachment point of the egg strand, within the interior of the mass, and in close contact with the substrate. This was attributed to the inability of the embryos to respire adequately and eliminate metabolic wastes. Biofouling did not strongly influence embryo mortality, but colonisation occurred in areas conducive to growth, photosynthesis, and respiration indicating ‘healthy’ regions within the mass.     

Reproductive behavior and spawning success of female Amblyglyphidodon leucogaster (Pisces: Pomacentridae) from the Red Sea

The reproductive behavior of female whitebelly damselfish, Amblyglyphidodon leucogaster, was investigated in the Gulf of Aqaba, Red Sea over two breeding seasons. Females were promiscuous, mating with 7–10 different males throughout the season. Females lay eggs in distinct batches, defined as the total number of eggs laid in a day. Generally females deposit a batch of eggs with one male (87.2%) and are capable of laying a new batch every other day. Egg batch size averaged 4009 eggs and females laid from 2 to 22 egg batches per season. The variation in spawning success was not correlated to body size. Females preferred to deposit eggs in nests that already contained early stage eggs (0–2 days old). Within a nest, females chose to lay eggs contiguous to the youngest egg batch, regardless if the nest contained either a single batch or multiple batches of different ages. Female within-nest spawning patterns appear to be a consequence of between nest preferences for nests with young eggs. It is proposed that the strong within-nest preference is a consequence of mate selection where females may use new egg batches as a visual cue as part of a copying style. Such a style may reduce the risk of predation and increase feeding opportunities, because less time is expended in mate selection, which would provide additional resources for egg production and ultimately increase female spawning success over the breeding season. This revised version was published online in July 2006 with corrections to the Cover Date.     

Ontogeny of squid mantle function: changes in the mechanics of escape-jet locomotion in the oval squid,Sepioteuthis lessoniana lesson, 1830

In Sepioteuthis lessoniana, the oval squid, ontogenetic changes in the kinematics of the mantle during escape-jet locomotion imply a decline in the relative mass flux of the escape jet and may affect the peak weight-specific thrust of the escape jet. To examine the relationship between ontogenetic changes in the kinematics of the mantle and the thrust generated during the escape jet, we simultaneously measured the peak thrust and the kinematics of the mantle of squid tethered to a force transducer. We tested an ontogenetic series of S. lessoniana that ranged in size from 5 to 40 mm dorsal mantle length (DML). In newly hatched squids, thrust peaked 40 ms after the start of the escape jet and reached a maximum of between 0.10 mN and 0.80 mN. In the largest animals, thrust peaked 70 ms after the start of the escape jet and reached a maximum of between 18 mN and 110 mN. Peak thrust was normalized by the wet weight of the squid and also by the cross-sectional area of the circumferential muscle that provides power for the escape jet. The weight-specific peak thrust of the escape jet averaged 0.36 in newly hatched squid and increased significantly to an average of 1.5 in the largest squids measured (P < 0.01). The thrust per unit area of circumferential muscle averaged 0.25 mN/mm(2) in hatchlings and increased significantly to an average of 1.4 mN/mm(2) in the largest animals tested (P < 0.01). The impulse of the escape jet was also lowest in newly hatched individuals (1.3 mN. s) and increased significantly to 1000 mN. s in the largest squids measured (P < 0.01). These ontogenetic changes in the mechanics of the escape jet suggest (1) that propulsion efficiency of the exhalant phase of the jet is highest in hatchlings, and (2) that the mechanics of the circumferential muscles of the mantle change during growth.     

Determining the daily spawning cycle of the chokka squid, Loligo reynaudii off the South African Coast

The study investigated the duration and peak of the daily spawning cycle of the chokka squid Loligo reynaudii, and the possibility of estimating the spawning frequency by means of classification and grouping (by age) of post-ovulatory follicles. Post-ovulatory follicles were classified into three types: new, intermediate or old, based in part on the degree of resorption of the follicles determined from histological sections. Ovulation in this species appears to be a rapid process, seen by the occurrence of both mature oocytes and post-ovulatory follicles in partially spent ovaries. Results suggest that chokka squid present a diel timing of spawning, with successive periods of egg accumulation at night, interrupted by periods of active spawning during the day. The ovarian cycle for this species appears typical of serial spawners. During the spawning phase ovaries go through ripe, partially spent and recovering stages by undergoing a process of maturation, ovulation and redeveloping, where a new batch of advanced oocytes is recruited. The cycle typically appears to last between 24 and 36 h, and may depend on a number of factors such as environmental conditions and the presence of predators. The short-lived (about 14 h) nature of the post-ovulatory follicles precludes the calculation of the exact spawning interval and does not allow us to accurately predict the spawning frequency. Together with the degree of oviduct fullness they do however give a good indication of imminent or recent spawning.     

Terminal spawning in the deepwater squid Moroteuthis ingens (Cephalopoda: Onychoteuthidae)

Maturation in the onychoteuthid squid Moroteuthis ingens was found to be irreversible, with death following shortly after sexual maturation and spawning. Both males and females were found with spent gonads. The ovary reaches very large sizes in mature females and probably prevents feeding by constricting the caecum. There was also a marked difference in the tissue integrity between immature females and females which had reached sexual maturity. Mature and spent females showed advanced tissue breakdown with individuals having a thin mantle wall with an inelastic, gelatinous appearance. Histological examination of the mantle wall revealed that the tissue breakdown was due to a drastic histolysis of muscle tissue and, to a lesser extent, collagen fibres. Mature males also showed some tissue breakdown and loss of muscle fibres but this was not as dramatic as in the females. These features are considered in relation to processes contributing to terminal maturation in M. ingens.     

Chromatophore activity during natural pattern expression by the squid Sepioteuthis lessoniana: contributions of miniature oscillation

Squid can rapidly change the chromatic patterns on their body. The patterns are created by the expansion and retraction of chromatophores. The chromatophore consists of a central pigment-containing cell surrounded by radial muscles that are controlled by motor neurons located in the central nervous system (CNS). In this study we used semi-intact squid (Sepioteuthis lessoniana) displaying centrally controlled natural patterns to analyze spatial and temporal activities of chromatophores located on the dorsal mantle skin. We found that chromatophores oscillated with miniature expansions/retractions at various frequencies, even when the chromatic patterns appear macroscopically stable. The frequencies of this miniature oscillation differed between "feature" and "background" areas of chromatic patterns. Higher frequencies occurred in feature areas, whereas lower frequencies were detected in background areas. We also observed synchronization of the oscillation during chromatic pattern expression. The expansion size of chromatophores oscillating at high frequency correlated with the number of synchronized chromatophores but not the oscillation frequency. Miniature oscillations were not observed in denervated chromatophores. These results suggest that miniature oscillations of chromatophores are driven by motor neuronal activities in the CNS and that frequency and synchrony of this oscillation determine the chromatic pattern and the expansion size, respectively.     

FMRFamide elicits chromatophore expansion and retraction depending on its type and development in the squid, Sepioteuthis lessoniana

To examine chromatophore control by FMRFamide-related peptide (FaRP), we investigated the pharmacological effect of FMRFamide on the chromatophores and the FMRFamide-immunoreactivity of nerves surrounding the muscles in the coastal squid, Sepioteuthis lessoniana. Applications of FMRFamide elicited expansion of black chromatophores and retraction of yellow chromatophores in the adult squid. FMRFamide-immunoreactive terminals were distributed along black chromatophore muscles but were not observed around the yellow ones. This means that FMRFamide functions differently for each of the two types of chromatophores in the adult squid. Moreover, the pharmacological effect of FMRFamide on the black chromatophores differed between adults and hatchlings; application of FMRFamide retracted black chromatophores in hatchlings but not in adults. These results indicate that certain squid species have an FaRP system for controlling the chromatophores in their skin and that the system changes during development.     

A new species, Trichiurus australis (Perciformes: Trichiuridae), from Australia

Trichiurus australis, a new trichiurid fish, is described on the basis of four specimens collected off Burnett River mouth, Pialba and Thursday Island, Queensland, Australia. The new species strongly resembles Trichiurus brevis Wang and You, 1992 (distributed in the South China Sea), T. russelli Dutt and Thankam, 1967 (the northwestern Bay of Bengal and the northern Gulf of Thailand), and T. nickolensis Burhanuddin and Iwatsuki, 2003 (northwestern Australia) in having the highest point of the supraoccipital crest situated directly above the posterior margin of the eye and being relatively small in size (less than ca. 700mm in total length: TL). It differs from those three species in having spinescent gill rakers almost with 2 equally long cusps [vs. (1, rarely 2 but 1 of them clearly shorter than another)], posterior caudal peduncle vertebrae bearing neural spines (vs. neural spines absent), longer caudal peduncle length (mean 16% TL vs. 6% in T. brevis, 8% in T. nickolensis, and 8% in T. russelli), and shorter precaudal length (83% vs. 93%, 91%, and 91%), preanal length (26% vs. 33%, 31%, and 35%), and head length (9% vs. 12%, 11%, and 13%). We have tentatively classified this new species under the Trichiurus russelli complex because the foregoing diagnostic characters are identical to this species complex except for the presence of neural spines in the posterior caudal vertebrae.     

Ontogenetic changes in fibrous connective tissue organization in the oval squid, Sepioteuthis lessoniana Lesson, 1830.

Ontogenetic changes in the organization and volume fraction of collagenous connective tissues were examined in the mantle of Sepioteuthis lessoniana, the oval squid. Outer tunic fiber angle (the angle of a tunic collagen fiber relative to the long axis of the squid) decreased from 33.5 degrees in newly hatched animals to 17.7 degrees in the largest animals studied. The arrangement of intramuscular collagen fiber systems 1 (IM-1) and 2 (IM-2) also changed significantly during ontogeny. Because of the oblique trajectory of the IM-1 collagen fibers, two fiber angles were needed to describe their organization: (1) IM-1(SAG), the angle of an IM-1 collagen fiber relative to the squid's long axis when viewed from a sagittal plane and (2) IM-1(TAN), the angle of an IM-1 collagen fiber relative to the squid's long axis when viewed from a plane tangential to the outer curvature of the mantle. The sagittal component (IM-1(SAG)) of the IM-1 collagen fiber angle was lowest in hatchling squid (32.7 degrees ) and increased exponentially during growth to 43 degrees in squid with a dorsal mantle length (DML) of 15 mm. In squid larger than 15 mm DML, IM-1(SAG) fiber angle did not change. The tangential component (IM-1(TAN)) of IM-1 collagen fiber angle was highest in hatchling squid (39 degrees ) and decreased to 32 degrees in the largest squid examined. IM-2 collagen fiber angle (the angle of an IM-2 collagen fiber relative to the outer surface of the mantle) was lowest in hatchling squid (34.6 degrees ) and increased exponentially to about 50 degrees in 15-mm DML animals. In squid larger than 15 mm DML, IM-2 fiber angle increased slightly with size. The volume fraction of collagen in IM-1 and IM-2 increased 68 and 36 times, respectively, during growth. The ontogenetic changes in the organization of collagen fibers in the outer tunic, IM-1, and IM-2 may lead to ontogenetic differences in the kinematics of mantle movement and in elastic energy storage during jet locomotion.     

Reproductive variance in planar spawning Chromodoris species (Mollusca: Nudibranchia)

Although variance in life history parameters is well known from comparisons among broad phylogenetic groups of marine invertebrates, there is still an outstanding need to increase empirical studies that compare closely related species. If the species under study share a recent common ancestor and developmental strategy, there is an opportunity to contrast maternal investment against interspecific variation while controlling for evolutionary distance. Furthermore, when these species co-occur, it allows for exploration of potential character displacement. We examined egg size and other factors related to reproduction in four closely related species, co-occurring nudibranchs belonging to the monophyletic Chromodoris planar spawning clade. The duration of oviposition appeared to be conserved and may be phylogenetically constrained in these four co-occurring species. In contrast, egg size differed significantly among species, but was not influenced by parental body length or position within the egg mass for any of the species. The number of egg mass whorls also varied, but did not correspond to the interspecific differences in parental body length. These results suggest that some significant differences exist among these sympatric Chromodoris species that may be candidate traits for character displacement. These characters would need to be re-measured in geographic areas where the studied Chromodoris species do not co-occur.     

Reproduction and larval biology in tunas,and the importance of restricted area spawning grounds

Tunas show a wide variety of life history strategies, spatial distributions and migratory behaviors, yet they share a common trait of spawning only in tropical and sub-tropical regions. The warm-water tuna species generally show significant overlap between spawning and feeding grounds, and longer spawning seasons of several months to near year-round. In contrast, the cool-water bluefin tunas migrate long distances between feeding and spawning grounds, and may spawn over periods as short as 2 months. Here, we examine the spatial distributions of tuna larvae in the world’s oceans, and examine interspecific differences in the light of adult behaviors and larval ecology. We discuss the links between larval tuna and their oceanographic environments and relate these to current knowledge of larval growth, feeding and trophodynamics, with a focus on the better-studied bluefin tunas. We show that larval tunas have moderate to fast growth rates and selective feeding habits, and thus appear to be adapted for survival in warm, oligotrophic seas. We also examine the challenges of sustainably managing species which migrate across multiple management boundaries to reach spatio-temporally restricted spawning grounds and discuss the previous and future anthropogenic impacts on tuna spawning areas.     

Ontogenetic changes in mantle kinematics during escape-jet locomotion in the oval squid, Sepioteuthis lessoniana Lesson, 1830.

We investigated the kinematics of mantle movement during escape jet behavior in an ontogenetic series of Sepioteuthis lessoniana, the oval squid. Changes in mantle diameter during the jet were measured from digitized S-VHS video fields of tethered animals that ranged in age from hatchlings to 9 weeks. The amplitude of both mantle contraction and mantle hyperinflation (expressed as percent change from the resting mantle diameter) during an escape jet was significantly greater in hatchlings than in older, larger squid (P < 0.05). The maximum amplitude of mantle contraction during the escape jet decreased from an average of -40% in hatchlings to -30% in the largest animals studied. The maximum amplitude of mantle hyperinflation decreased from an average of 18% in hatchlings to 9% in the largest squid examined. In addition, the maximum rate of mantle contraction decreased significantly during ontogeny (P < 0.05), from a maximum of 8.6 mantle circumference lengths per second (L/s) in hatchlings to 3.8 L/s in the largest animals studied. The ontogenetic changes in the mantle kinematics of the escape jet occurred concomitantly with changes in the organization of collagenous connective tissue fiber networks in the mantle. The alteration in mantle kinematics during growth may result in proportionately greater mass flux during the escape jet in newly hatched squid than in larger animals.