Continuous spawning in the cirrate octopods Opisthoteuthis agassizii and O. vossi: features of sexual maturation defining a reproductive strategy in cephalopods

The reproductive strategy of the cirrate octopods Opisthoteuthis agassizii and O. vossi (collected off Namibia from 1988 to 1990) was analyzed. Ovarian oocyte size frequency analysis for both species revealed continuous egg production over the entire adult life span. Mature eggs were stored in the single oviducal gland and distal oviduct, but oviducal gland fullness was not related to body size (p>0.2). All O. agassizii male specimens from 95 to 5400 g total weight were sexually mature, as were all females from 190 to 1650 g, indicating that considerable growth takes place after the onset of sexual maturity. Continuous spawning is defined as a single, extended and continuous period of egg maturation and spawning. This model of reproductive strategy is previously unreported in cephalopods. All O. vossi male specimens from 750 to 3050 g total weight, and females from 800 to 1300 g, were sexually mature. Mature males and females of both species were collected in all seasons of the year. The adaptation of cirrate octopods to non-scasonal deep-sea environments is considered. The sexual maturity characteristics of males were analyzed, and examination of the spermatophore revealed opercular structures previously unreported in cephalopods. For females, the micropyle of the eggs are described and the mineral analysis of the egg shell disclosed that sulphur was the major element present.     

Trophic and reproductive biochemistry of a deep-sea gelatinous octopus, Opisthoteuthis calypso

The present study provides a comprehensive analysis of the biochemical composition (proximate composition, amino acids and fatty acids in gonad, digestive gland and muscle) of the cirrate octopod, Opisthoteuthis calypso, collected off the coast of Portugal. Protein and energy contents of O. calypso were significantly lower (p < 0.05) than those observed for coastal octopods, suggesting that the gelatinous musculature of the cirrate octopus may not be a direct consequence of food limitation, but rather associated with reduced selective pressure for strong swimming ability in the deep sea. Moreover, principal component analyses of fatty acid (FA) data clearly separated coastal octopods from O. calypso. Saturated FA biomarkers (namely 16:0, 17:0, 18:0) and monounsaturated FA 22:1 suggest a higher phytodetrital influence on diet of O. calypso. A clear separation between groups was also observed with amino acid (AA) data. However, essential AA and non-essential AA designations (based on the flow of carbon through biochemical systems) do not necessarily provide an accurate picture of the origins of amino nitrogen in the deep-sea environment. Consequently, the interpretation of the present data is a challenging task but opens a new window of opportunity to unravel new trophic biomarkers in the deep sea.     

Buoyancy in Porcellana platycheles

Intermoult Porcellana platycheles (Pennant) are slow moving, crawling crabs, incapable of swimming upwards from the substrate. Newly moulted porcelain crabs, however, are capable of effective upward swimming. An increased efficiency of tail-fan propulsion in newly moulted crabs was thought unlikely, and it was suspected that newly moulted P. platycheles were more buoyant than intermoult crabs. Results are presented to show that this is the case, and calculations were performed which indicate that fluid uptake at moult is insufficient to produce the observed increase in buoyancy. It is concluded that loss of heavy minerals and scleroprotein at moult cause the increased buoyancy. The adaptive significance of the enhanced swimming ability in newly moulted crabs is thought to lie in an improved ability to escape predation by intertidal fish and crabs.     

Sequence divergence of mitochondrial DNA indicates cryptic biodiversity in Octopus vulgaris and supports the taxonomic distinctiveness of Octopus mimus (Cephalopoda: Octopodidae)

DNA sequence diversity of octopods was investigated using the mitochondrial cytochrome oxidase III gene (mtCOIII). DNA was obtained from ethanol- or formalin-fixed tissue of 15 specimens belonging to Octopus mimus Gould, 1852, Octopus vulgaris Cuvier, 1797 and Scaeurgus unicirrhus d'Orbigny, 1840, from coastal waters of the Mediterranean (France), the southwestern Atlantic (Brazil), the Caribbean (Costa Rica) and the southeastern/tropical Pacific (north Chile/Costa Rica). A 612 bp fragment of the mtCOIII gene was sequenced and aligned to the orthologous sequences available from northeastern Pacific Octopus species. Possible phylogenetic relationships were reconstructed. The mtCOIII gene tree revealed two main clusters, one comprised O. rubescens, O. dofleini and O. californicus, while the other included all O. vulgaris specimens, O. bimaculatus, O. bimaculoides and O. mimus. With one exception all South American haplotypes including O. bimaculatus and O. bimaculoides appeared as the sister group of the Mediterranean haplotype of O. vulgaris, indicating that most of the South American O. vulgaris specimens investigated would not belong to the recently redescribed species O. vulgaris restricted to the Mediterranean and eastern Atlantic. The treatment of O. mimus as a species distinct from Mediterranean O. vulgaris is supported by a high nucleotide divergence of 12.7%. Based on the mtCOIII gene tree the existence of cryptic species among O. vulgaris-like octopods is suggested. Received: 15 January 1999 / Accepted: 15 October 1999     

Cementation vs mobility: development of a cemented byssus and flexible mobility in Anomia chinensis

The mobility of Anomia chinensis Philippi was studied in relation to its byssal development stage. This species shows high mobility even after it begins cementation in its post-larval stage. Juveniles develop a calcified byssus in the post-larval stage and cement to substrata. However, juveniles up to about 10 mm in shell length can relocate by repeating a sequence of formation of the calcified byssus, abandonment of it, locomotion by crawling, reattachment, and recementation. Juvenile anomiids up to 25 mm in shell length also can move, without breaking their byssal attachments, by shifting the center of byssal calcification dorsally. Even an adult can change its orientation by forming a twisted byssus. These possible methods of movement are closely related to five stages of byssal development. Anomiids can use this mobility to seek a preferable position for attachment after initial cementation, or to adjust their orientation, and thus promote higher survivorship. Received: 18 August 1997 / Accepted: 19 July 1998     

Fecundity and reproductive strategies in deep-sea incirrate octopuses (Cephalopoda: Octopoda)

Coleoid cephalopods show flexibility in their reproductive strategies or mode of spawning, which can range from simultaneous terminal spawning over a short period at the end of the animal’s life to continuous spawning over a long period of the animal’s life. Although a simultaneous terminal spawning strategy is typical of shallow water temperate octopuses, it is not known whether deep-sea octopods would have the same reproductive strategy. The reproductive strategies and fecundity were investigated in nine species of deep-sea incirrate octopuses: Bathypolypus arcticus, Bathypolypus bairdii, Bathypolypus ergasticus, Bathypolypus sponsalis, Bathypolypus valdiviae, Benthoctopus levis, Benthoctopus normani, Benthoctopus sp., and Graneledone verrucosa (total n = 85). Egg-length frequency graphs and multivariate analysis (principal components analysis) suggest that B. sponsalis has a synchronous ovulation pattern and therefore a simultaneous terminal spawning strategy. Although a simultaneous terminal spawning strategy is most likely for B. levis and B. normani, the egg-length frequency graphs and multivariate analysis also suggest a greater variation in egg-lengths which could lead to spawning over an extended period.     

Reduced take-off ability in robins (Erithacus rubecula) due to migratory fuel load

Recent studies have shown that large fuel loads in small birds impair flying ability. This is the first study to show how migratory fuel load affects flying ability, such as velocity and height gained at take-off in a predator escape situation, in a medium-distance migrant, and whether they adjust their take-off according to predator attack angle. First-year robins (Erithacus rubecula) were subjected to simulated attacks from a model merlin (Falco columbarius), and take-off velocity and angle were analysed. Robins with a wing load of 0.19 g cm⁻² took off at a 39% lower angle than robins with a wing load of 0.13 g cm⁻², while velocity remained unaffected. The robins did not adjust their angle of ascent in accordance with the predator's angle of attack. Since many predators rely on surprise attacks, a difference in flight ability due to varying fuel loads found in migrating robins can be important for birds' chances of survival when actually attacked. Received: 28 October 1998 / Received in revised form: 12 January 1999 / Accepted: 30 January 1999     

Flipper beat frequency and amplitude changes in diving green turtles, <Emphasis Type="Italic">Chelonia mydas</Emphasis>

Using a turtle-borne camera system, changing flipper beat frequency and amplitude were measured in five diving green turtles (Chelonia mydas Linnaeus 1758) in the Bahía de los Angeles, Mexico (28°58′N, 113°33′W). These observations were made between June and August 2002. Turtles worked hardest (i.e., had the highest flipper beat frequency and amplitude) at the start of descents when positive buoyancy is predicted to oppose their forward motion. During the later part of descents, turtles worked less hard in line with opposing buoyancy forces being reduced. For example, flipper beat frequency declined from about 60–80 beats min⁻¹ at the start of descent to around 25–40 beats min⁻¹ after 30 s of the descent. At the start of ascents the flipper beat frequency was around 30 beats min⁻¹, lower than on descent, and declined as the ascent progressed with often passive gliding for the final few meters to the surface. This pattern of effort during diving appears to apply across a range of marine reptiles, birds and mammals suggesting that graded effort during descent and ascent is an optimum solution to minimising the cost of transport during diving.     

Association of Syscenus infelix (Crustacea: Isopoda: Aegidae) with benthopelagic rattail fishes, Nezumia spp. (Macrouridae), along the western North Atlantic continental slope

During submersible surveys along the continental slope (summers of 1991 and 1992, 184–847 m) between False Cape, Virginia, and Cape Hatteras, North Carolina, USA, we observed the aegid isopod, Syscenus infelix Harger, attached to the macrourid Nezumia bairdii (Goode and Bean). This is the first report of S. infelix attached to fishes in the western North Atlantic. The association of this blind isopod with its host appears species specific. The large, conspicuous isopod always attached to a fish in the same location, the dorsal midline, immediately behind the first dorsal fin. Attachment appears to be long term, with the isopod forming a characteristic scar consisting of a distinct discolored oval depression with seven small, dark impressions that coalesce as the fish grows. Only one S. infelix was found on each host fish. The isopod occurred on 23.7% of N. bairdii observed from submersible on the middle continental slope off Virginia and North Carolina, compared with 16.6% of 1236 museum specimens of the same species (based on inspection for scars) collected at latitudes 26°–64°N. Prevalence of the fish–isopod association was not correlated with depth or latitude. We also found identical scars on preserved specimens of N. aequalis (2.6% of 660 specimens), N. sclerorhynchus (1.2% of 86 specimens), and N. suilla (14.3% of 7 specimens), mostly from areas outside the range of N. bairdii. No scars were found on museum specimens of N. atlantica (n=27), N. cyrano (n=57), or N. longebarbata (n=7). The low incidence of isopod attachment on these species suggests that N. bairdii is the preferred host. Infestation by the isopod appears to result in erosion of host fish scales and tissue. We propose that S. infelix is an obligate associate of its host fish and should be considered parasitic. Received: 9 June 2000 / Accepted: 21 October 2000     

Effect of microbial films on settlement of bryozoan larvae (Bugula simplex,B. stolonifera and B. turrita)

The influence of microbial films on larval settlement was studied for sympatric species of cellularioid cheilostome bryozoans (Bugula simplex, B. stolonifera, and B. turrita). Settlement experiments were conducted in the laboratory in 1981 with and without a choice of filmed or unfilmed substrates. When given a choice of substrates, the larvae of all three bryozoan species significantly preferred the filmed substrate. When not given a choice of substrates, both B. simplex and B. turrita larvae tended not to settle on the unfilmed substrate, while the filmed and unfilmed substrates were equally attractive to B. stolonifera. Microbial films enhanced the settlement of all three species.     

Reproductive biology in two species of deep-sea squids

Deep-sea squids, Moroteuthis ingens and Gonatus antarcticus, were collected in the slope waters off the Falkland Islands and their reproductive systems preserved and investigated onshore. Changes in oocyte length-frequencies at maturation and spawning, and their fecundity were studied. These squids, as well as many other species, are characterised by a synchronous oocyte growth and ovulation. Oviducts are not used for ripe egg accumulation and consequently the universal scale of Lipinski (1979) cannot be applied to assign female maturity. M. ingens spawns near the bottom; its fecundity is 168–297 thousand eggs. Maximum egg size is 1.8–2.7 mm. G. antarcticus spawns midwater; its fecundity is 10–25 thousand eggs. Egg size is 3.2–3.3 mm. In M. ingens spawning takes place in the austral autumn and winter, in G. antarcticus—in austral winter. Our data and the literature data show that the so-called “synchronous ovulation” probably occurs in all deepwater squids. This pattern is very rare among fish, but is quite common among benthic octopods that brood their egg masses.     

Early life history traits of the gold-eye rockfish, Sebastes thompsoni, in relation to successful utilization of drifting seaweed

The gold-eye rockfish Sebastes thompsoni lives in association with drifting seaweed during the early developmental period. Early life history traits of the gold-eye rockfish were studied by means of otolith analysis of larvae and juveniles associated and unassociated with drifting seaweed in the waters off southern Sanriku, Tohoku, northern Japan, from 1993 to 1995.␣Daily growth increments and the existence of an extrusion check were confirmed by both a series of tetracycline marking experiments and an analysis of the relation between the number of increments and the number of days after extrusion. Larvae and juveniles associated with drifting seaweed occurred from late May to late July, with a peak in late June, showing the same seasonal fluctuation in abundance as drifting seaweed. Larvae seemed to associate with drifting seaweed after aggregation to surface slick areas. Body size at the commencement and the end of the seaweed-associated period was estimated at 18 and 50 mm total length, respectively. Growth rates were low until the beginning of the association with seaweed; thereafter they became high but declined towards the end of seaweed-associated life. Larvae and juveniles born later grew faster than those born earlier. This tendency was detected both before and during the association with drifting seaweed. Although the season of abundant drifting seaweed was about 2␣months shorter, and occurred 2 months later, than the extrusion season of the gold-eye rockfish from early February to early June, this seemed to be compensated by the growth variation dependent on birth month. Early life history traits of the gold-eye rockfish are discussed in relation to the successful utilization of drifting seaweed. Received: 3 August 1997 / Accepted: 24 March 1998     

Buoyancy of natural populations of marine phytoplankton

Buoyancy of natural populations of marine phytoplankton was studied in a fjord in western Norway during the diatom bloom and in autumn. The study was carried out under approximate in situ conditions by means of an apparatus described in the paper. During the spring bloom, positive buoyancy was observed only once. Sinking rates of individual fractions ranged from 0 to more than 9 m day^-1, and the mean sinking rates of the total chlorophyll content from 0 to at least 2. 2 m day^-1. The highest rates occurred in the post-bloom period, while sinking appeared negligible from the onset of the bloom up to its culmination. In autumn, the population was dominated by small, flagellated cells. Positive buoyancy, or upward migration, was then observed in two out of three experiments.     

Experimental analysis of the contribution of swimming and drifting to the displacement of reef fish larvae

The extent to which behaviour affects the dispersal of pelagic larvae in reef fishes has been a topic of major discussion among marine ecologists. Here, we experimentally quantified the extent to which the displacement of late-stage larvae of Abudefduf saxatilis is due to active movement (i.e. swimming) and drifting. We consider drifting as the component of larval displacement accounted for by the current. Drifting was quantified by comparing larval displacement to the displacement of passive particles in an extended flow chamber that gave larvae the free choice of swimming (i.e. swim with or against the current or not swim at all). We also determine whether drifting results from currents exceeding larval swimming capabilities or from the behavioural choice of larvae of not to swim against adverse currents. To do this, we compare the speeds of larval swimming in the extended flow chamber to those obtained in a smaller chamber in which larvae are behaviourally forced to swim due to space constraints and a retaining fence (most available data on larval swimming is based on this sort of chamber). Within the extended chamber, larvae tended to face the current and swim slower than it. This resulted in a net displacement increasingly determined by drifting. We also found that in the extended chamber, larvae swam at speeds between one and six times slower than the speeds they achieved in the “behaviourally modifying” smaller chamber. This suggests that the net displacement in the extended chamber was in part due to the behavioural choice of the larvae of not to swim. The importance of this “behavioural drifting” is discussed in terms of energy savings required for successful completion of the larval period and post-settlement survival. The idea that larvae may modulate their swimming behaviour raises caution for the use of published data regarding swimming capabilities of reef fish larvae when assessing the extent to which these fish actively affect their dispersal.     

Dispersal mechanisms in amphipods: a case study of <Emphasis Type="Italic">Jassa herdmani</Emphasis> (Crustacea,Amphipoda) in the North Sea

Jassa herdmani (Walker 1893), a tube-building amphipod typical of hard substrates, was found in large densities on shipwrecks from the Belgian part of the North Sea, in association with the hydrozoan Tubularia indivisa. In this area, shipwrecks only represent the source of hard substrates in an environment dominated by soft sediments. Nevertheless, the long-distance dispersal of Jassa species has never been investigated. Therefore, we tested the hypothesis of dispersal with currents, by investigating the behaviour of J. herdmani in the laboratory. Size distribution revealed that newly released juveniles (<1 mm) predominated throughout the year and the lower frequency of the size class 1–3 mm indicates a high mortality or a dispersal at this life stage. Individuals of J. herdmani may initiate actively the transport by tidal or surface currents by swimming to the surface of the water or by floating at the surface, as suggested by behaviours noted in the laboratory. The reaction of the amphipods to a current was investigated in the laboratory. We could not detect any sexual function associated with drifting and there was also no preponderance of a size class in the drifting individuals. When testing different substrates, we observed a significant influence of the substrate type on the frequency at which drifting occurred: J. herdmani showed a better adherence on T. indivisa compared to other substrates. Finally, the amphipod showed also a preference for its host compared to other substrates, which suggests a possible detection mechanism.     

Observations on underwater locomotion and flipper movement of the humpback whale Megaptera novaeangliae

Humpback whales (Megaptera novaeangliae) were observed and photographed during winter, 1974 near the West Indies and summer, 1976 near Newfoundland. Prior investigations on whale locomotion based on anatomical inference or surface observations did not describe any specific underwater use of the humpbacks' uniquely long flippers. Our results show that humpbacks achieve a high degree of maneuverability underwater through the active coordinated use of their flippers. Three-dimensional active movements including protraction-retraction, abduction-adduction, and lateral-medial rotations, were observed. The whales could move themselves in all directions independent from fluke-generated forward motion by thrusting with their resilient flippers. At higher swimming speeds (>4 knots) the flippers were used to effect sharply executed ascending or descending banked turns. Analysis of filmed sequences from three adult-calf encounters indicated a reduction of the amplitude and frequency of tailbeats by the adults; calves made tailbeats 4 to 7 times more frequently than adults when maintaining the same speed. The tail and flukes are forcefully moved other than vertically during maneuvering. The significance of these findings is discussed in relation to our knowledge of larger whales.     

Impact of submersible pumps on Pb constituents in residential wells

Dissolved Pb in 51 domestic wells screened from 18 to 48 m in glacial tills and outwash deposits were examined in conjunction with the characteristics of their corresponding submersible pump. Pb concentrations, ranging from 0.8 to 24.9 μg l⁻¹, entering residential water supplies were measured during 2001–2004 in the Royal watershed, Maine. Principal component analyses assisted the weighting of pump age, well screen depth, and draw time variables. Preliminary Pb sequestration significance in the boreholes was predicted from geochemical speciation and synchrotron XAS analyses. Nascent ²⁰⁷Pb/²⁰⁶Pb and ²⁰⁸Pb/²⁰⁶Pb isotope analyses assisted the discrimination of possibly leached Pb from submersible pump materials among geogenic sources.     

Crossing the frontier: vertical transit rates of deep diving cormorants reveal depth zone of neutral buoyancy

Knowing the depth zone of neutral buoyancy of divers is important because buoyancy can determine how animals manage their energy budget. In this study, we estimate the depth zone of neutral buoyancy of free-ranging cormorants for the first time, using time-depth recorders. We discovered that vertical ascent rates of 12 Crozet and 15 Kerguelen diving blue-eyed shags (respectively Phalacrocorax melanogenis and P. verrucosus) slowed down considerably at the 50–60 m depth zone. We suggest this was due to birds trying to reach the surface from that point upwards using reduced locomotor activity because the force of buoyancy becomes greater than the force of gravity at that depth. The results show a shift of this depth zone in relation to maximum targeted dive depth, suggesting cormorants may control buoyancy through respiratory air volume adjustment. Interestingly, 60 m is close to the maximum depth zone reached by these two species during dives lasting 4 min, their estimated behavioural aerobic dive limit. This suggests that the decision to swim deeper has a direct consequence on the energy budget, with time spent recovering at the surface (time thus lost to foraging) strongly increasing relative to the preceding time of submergence. Resources found in deeper waters must be of sufficient quantity or quality to justify crossing the frontier of physical neutral buoyancy.     

Dispersal potential of invasive algae: the determinants of buoyancy in <Emphasis Type="Italic">Codium fragile</Emphasis> ssp. <Emphasis Type="Italic">fragile</Emphasis>

The capacity for long-distance dispersal is an important factor in determining the spread of invasive species. For algae, positive buoyancy generally is correlated with increased dispersal potential, and the light environment has been previously identified as a possible determinant of buoyancy in several species. We examined the effect of light intensity on the buoyancy of fragments of the invasive green alga Codium fragile ssp. fragile. Under natural and controlled conditions, the buoyancy of samples taken from the thallus tip was higher than those from near the holdfast. Both laboratory and field experiments also showed that buoyancy was dynamic and switched from positive to negative under reduced light intensity, but this change required several days. We also observed seasonal changes in buoyancy, presumably due to natural variations in light intensity, with the buoyancy of fragments washed up on the shore highest in mid-summer. These results show that buoyancy is a dynamic property of the C. fragile ssp. fragile thallus and suggest that buoyant fragments contribute to long-range dispersal and accelerated regional spread of this invader. This finding suggests that dispersal is more likely during conditions of high light intensity and illustrates the need to understand how variations in the natural environment can affect the dispersal potential of invasive species.     

Life on glass houses: sponge stalk communities in the deep sea

Photographs of the deep-sea floor often show organisms attached to biogenic structures that protrude from the soft bottom. In particular, the stalks of glass sponges (hexactinellids) provide hard substrata and act as habitat islands for deep-sea fauna. The primary objectives of this study were to determine the abundance of glass sponge “stalks” at an abyssal station in the NE Pacific, to identify the fauna associated with stalks, and to compare the distribution patterns of epifaunal taxa both horizontally and vertically. Densities of stalks and large epifauna were estimated from analysis of ∼9 km of photographic transects taken in 1994–1995 at station M (34°45′N; 123°00′W; 4,100 m depth) off California, USA. At least 87% of the stalks were the spicule columns of live or dead hexactinellids in the genus Hyalonema (Gray, 1832). Stalks appeared to be distributed randomly across the sea floor (density: 0.13 stalks m⁻²). A colonial zoanthid, Epizoanthus stellaris (Hertwig, 1888), inhabited 20% of the stalks and was the most commonly observed epifaunal organism, followed by other suspension feeders that generally were situated at the top of the structures. Thirty-five stalk communities were collected in tube cores in 1994–1995 using the submersible “Alvin”. A total of 139 taxa was associated with these hard-substratum habitats (another five species were observed only in photographs). Although taxon richness was high, the species diversity of these communities was relatively low due to the dominance in percentage abundance of a foraminiferan, Cibicides lobatulus (Walker and Jakob, 1798), and a serpulid polychaete, Bathyvermilia sp. (Zibrowius, 1973). The relationship between number of taxa and surface area of the stalks yielded a slope (z-value) typical of islands with a low rate of immigration. Three-dimensional complexity created by branching epifauna on the stalks provided more surface area and a variety of cryptic microhabitats. Vertical zonation on the stalks appeared to be controlled by biological interactions among species, with solitary fauna and certain functional groups of colonial organisms restricted by sheet-like colonial organisms that appeared to be dominant space competitors. Received: 13 April 2000 / Accepted: 9 November 2000