ALBERT

Notes: Abstract Most studies of spatial patterns of invertebrates in soft sediments have concentrated on populations of individual species. Those that examined patterns in communities have tended to employ categorical analytical techniques. Using macrofaunal abundance data from van Veen grab samples collected 20 to 100 m apart in known spatial arrangements from Scottish sea-lochs, the relationships between patterns in macrobenthic species composition and distances between samples were explored using matrix correlations in a non-parametric framework. Using a simple definition of spatial structure, i.e. that intersample distances are monotonically correlated with intersample species-similarity, spatial structure at each of seven stations was assessed using non-parametric Mantel tests based on rank-correlations. Changes in community structure were positively correlated with distance at all sites in Loch Etive, on both current-swept muddy sands and soft deep muds. Different components of the macrobenthos contributed to spatial pattern at each site. Simple spatial structure was also detectable at a muddy-sand site in Loch Creran, but neither on soft mud, nor at the soft mud site in the Firth of Lorne. The concept of rank-correlograms was introduced. These were used to examine the extent and form of spatial structure in different components of the macrobenthos at each site. Relationships between similarity and distance were neither simple nor consistent. Results were compared to previous studies which used the same data, and it was concluded that studies carried out at a particular scale, or on a particular component of the benthos, are unlikely to be successful in predicting spatial relationships at other scales or for other components of the benthos. Correlational rather than categorical analyses are recommended for exploratory studies of spatial relationships in the benthos. Analyses of the spatial structure at these seven sea-loch sites suggests that by ensuring that samples are at least 40 m apart an investigator is unlikely to underestimate variability or otherwise invalidate statistical analyses based on the use of the samples as replicates. Spacing samples up to 100 m apart may increase variability estimates, further reducing the chance of concluding that a difference exists when one does not.

Notes: Abstract   Ophiocten gracilis is an ophiuroid found at bathyal depths in the North Atlantic Ocean. The adults show strong seasonal reproduction, with an ophiopluteus in the surface plankton. Settling postlarvae were collected in sediment traps moored at 1000 and 1400 m depth in the NE Atlantic during Julian Days 142 to 212 (May to July) in 1996. During this period, growth of postlarvae in the traps was linear and the diet consisted of phytodetritus and foraminifera. Experiments suggest that postlarvae sink at rates of up to 500 m d−1, although this may well be slower in the natural environment. The high fecundity, seasonality and high population density resulted in high fertilization success, and many of the offspring were advected outside the normal adult range, where they were able to settle but did not survive to adulthood.

Notes: Abstract Samples of the deep-sea spinulosan asteroid Hymenaster membranaceus Wyville Thomson were collected in a timeseries of 19 bottom trawls spanning the period April 1978 to October 1981 from a 2200 m-deep station in the northern Rockall Trough. The reproductive biology of this species was studied from histological sections of the gonad, and compared with that of H. gennaeus H. L. Clark which was collected in the same hauls. At first sexual development, oogonia develop in nests surrounded by small accessory cells. Previtellogenic oocytes remain in the periphery but, at maturity, oocytes ranging up to 1 100 μm fill the ovary. A variety of accessory cells pack the lumen and may be nutritive or degenerative. It appears that a small number of oocytes are spawned intermittently, but there is no evidence of overproduction and break-down of superfluous oocytes. A few large oocytes become senescent and undergo internal break-down, releasing periodic acid Schiff-positive material into the lumen. Size-frequencies of oocytes indicate that eggs may be spawned as a continuous slow release, and there is no evidence of reproductive synchrony between or within samples. On reaching maturity, males appear always to be ready to release spermatozoa. Spawning is probably stimulated by egg release during chance encounters with mature females. There is no evidence for brooding, and from the large size and yolky nature of the egg direct lecithotrophic development at or near the seabed is inferred. A limited histological study of H. gennaeus indicates that egg production is very similar, but the two species differ in the nature of the accessory cells and amorphous material filling the lumen.

Notes: Abstract Growth bands have been found in the calcitic vertebral arm ossicles of the commonly occurring deep-sea brittle starsOphiura ljungmani Wyville Thomson andOphiomusium lymani (Lyman) (Echinodermata: Ophiuroidea) trawled from the Rockall Trough (N.E. Atlantic) at 2 200 and 2 900 m depth from 1973 to 1982. InOphiura ljungmani, the study of ossicle microstructure by SEM shows that growth bands reflect differences in stereom porosity and surface relief, similar to that previously found amongst shallow-water brittle stars. The pattern inOphiomusium lymani was much less clear from the microstructure, but could be revealed by heating the ossicle to 450 °C and “clearing” in xylene. The bands showed up as fine, translucent rings separated by more opaque, wider zones, perhaps reflecting differences in organic material incorporated within the calcite. Both growth-banding patterns probably reflect an annual cycle in skeletal growth rate. On this assumption, the number and spacing of the banding in the two species indicates contrasting growth strategies. WithOphiura ljungmani, a rather regular annual growth increment and perhaps shorter lifespan (up to ca. 10 yr) thanOphiomusium lymani is indicated. The latter shows a relatively wide spacing of early bands, followed by tight clustering of the outermost bands corresponding to adult sizes. This growth pattern is characteristic of species “escaping” from predation by rapid growth to relatively large adult size. AdultO. lymani probably grow slowly, some perhaps reaching 20 yr of age. Growth curves were fitted to size-at-age corresponding to measurements of the size and ordering of growth bands. These corroborate age structure previously estimated from analysis of size frequencies in time series from the stations sampled in the present study. Such skeletal growth markers should be of value in analysis of the demography of deep-sea populations.

Notes: Abstract Although the taxonomy of deep-sea protobranch bivalves is becoming better known, relatively little information is available on their reproductive biology and whether or not populations show reproductive periodicities. We have examined the reproductive biology of three common sympatric species as part of a long-term time-series of samples taken from 2900 m in the Rockall Trough from 1973 to 1983. Malletia cuneata Jeffreys, 1876 produces a maximum of 30 oocytes at any one time and these grow to a maximum size of 240 μm. Maximum fecundity of Ledella pustulosa (Jeffreys, 1876) and Yoldiella jeffreysi (Hidalgo, 1877) is 174 and 360, respectively, and both species produce an egg of ∼ 120 μm in diameter. These data indicate lecithotrophic early development in L. pustulosa and Y. jeffreysi, but direct development in M. cuneata; however, evidence from the prodissoconch length of M. cuneata suggests lecithotrophic development. L. pustulosa and Y. jeffreysi also differ from M. cuneata in having a distinct reproductive cycle with spawnout in the winter months. Both the continuously breeding M. cuneata and the seasonally breeding L. pustulosa ingest diatoms, coccoliths and foraminiferans, but whereas the diet of M. cuneata appears to be constant throughout the year there is an apparent reduction in the feeding activity of L. pustulosa concomitant with the deposition of phytodetritus on the deep-sea bed.

Notes: Abstract Growth of the European edible sea urchin Echinus esculentus L. was studied in a population held for 2 yr in cages on the sea bed, after labelling with the skeletal growth marker tetracycline. The final position of the tetracycline tag on the genital plates agreed with an annual periodicity in natural growth zones; two such growth zones appeared beyond the position of the tag on the ground surface of the plate as light-reflecting bands separated by narrow dark lines in the middle layer. Individual and group (pooled data) growth parameters were estimated from the growth increment shown in the genital plate, whose lateral growth displayed a linear relationship to the diameter of the urchin test within the size range of these measurements. Von Bertalanffy growth parameters (asymptotic size and growth-rate function K) fitted to the growth increment on each individual were highly significantly correlated to those fitted to the natural growth lines, assuming an annual periodicity. The inferred growth pattern agrees well with conclusions based on H. B. Moore's growth-band data. The caged urchins can be assumed to have experienced exactly similar conditions, yet the growth curves fitted to individuals showed considerable variability. The good agreement between estimated growth function parameters of individuals obtained by the two methods indicate that this reflects real variability in growth between individuals that probably is genetically rather than environmentally determined. The growth of E. esculentus, and the adaptational significance of high growth variability in the population is briefly discussed.

Notes: Abstract Examination of the diet of two sympatric species of seastar, Bathybiaster vexillifer and Plutonaster bifrons from ∼2200 m depth in the Rockall Trough, NE Atlantic Ocean (∼57°18'N; 10°28'W), suggested that diet may determine the different reproductive patterns found between these two species. In the non-seasonally breeding B. vexillifier, the diet showed a high Shannon-Wiener prey diversity index, the dominant prey being the irregular echinoid Hemiaster expergitus together with a variety of prosobranch gastropods and protobranch bivalves. By contrast, the prey diversity in the seasonally breeding P. bifrons was significantly lower than that of B. vexillifer. In addition, organic carbon content in the sediment residue in the stomachs of P. bifrons displayed a seasonal cycle, while no such seasonality was detected in B. vexillifer. The stomachs of P. bifrons also contained a higher proportion of scavenged material, including the seasonally available remains of mesopelagic blue whiting. These data, together with “Bathysnap” (time-lapse camera) observations of feeding behaviour in both species, suggest that B. vexillifer is a predator feeding deep in the sediment, whereas P. bifrons feeds close to the sediment surface where it is affected by the seasonal availability of phytodetritus and fish carcasses.

Notes: Abstract Aspects of the reproduction and population biology of two elasipodid holothurians collected during the period 1973–1984 from the north-east Atlantic Ocean were examined. The reproductive biology of both the benthic species Peniagone azorica and the benthopelagic P. diaphana are similar. A primary oocyte forms from an oogonium and grows to about 100μm diameter, whereupon it undergoes vitellogenesis and increases to 300μm before being spawned. A high proportion of primary oocytes are not spawned and undergo a complicated breakdown process resulting in the formation of an amorphous sac in the ovary wall. It is possible that when an ovarian tubule is full of these sacs it atrophies and drops off, allowing other tubules to develop. The maximum egg size for both species suggests abbreviated larval development. In P. azorica, juveniles reach sexual maturity for the first time at about 30 mm length. The population structure suggests that the adults grow slowly, although it is possible that the observed unimodal distribution in size-frequencies results from one or several recruitments. Recruitment to the population is probably infrequent and may occur irregularly. Comparison of the population structure of closely spaced samples suggests a patchy distribution on the bottom. The benthopelagic lifestyle of adult P. diaphana suggests that the larvais also planktonic.

Notes: Abstract The reproductive biology of 5 species of echinothuriid (Phormosoma placenta, Calveriosoma hystrix, Araeosoma fenestrum, Sperosoma grimaldii and Hygrosoma petersii) and 2 species of cidarid (Cidaris cidaris and Poriocidaris purpurata) sea urchins from the deep sea (Rockall Trough) has been examined from samples collected during 1973–1983. In all species the gonads lie within the interambulacrum attached to aboral gonopores and when fully developed occupy most of the test not occupied by the gut or Aristotle's lantern. In all the species, initial oocyte development takes place along the germinal epithelium embedded in nutritive tissue. In all the echinothuriids and in Poriocidaris purpurata, the oocyte grows to ca. 200 to 450 μm, at which stage vitellogenesis begins. Oocyte growth continues until a maximum egg size of 1 100 to 1 500 μm is attained. In the echinothuriids, two types of nutritive tissue are found. In the carly stages of gametogenesis the oocyte is surrounded by well-structured periodic acid Schiff (PAS)-positive tissue. As the oocyte grows this tissue becomes vacuolated, suggesting that there is a transfer of nutriment to the developing oocyte. In Phormosoma placenta, unspawned oocytes are phagocytosed. There is no evidence of seasonality in any of the echinothuriid species or in Poriocidaris purpurata. Extrapolation with shallow-water echinothuriids suggests that larval development is lecithotrophic, omitting any planktotrophic phase. Of the species examined, only Cidaris cidaris has a reproductive strategy which produces a known larva, although the limited samples did not permit any determination of seasonality in this deep-sea population.

Notes: Abstract The stomachs of 1 165 individuals of the species Ophiacantha bidentata (Retzius), Ophiactis abyssicola (M. Sars), Ophiocten gracilis (O. Sars), Ophiura irrorata (Lyman), O. ljungmani (Lyman) and Ophiomusium lymani Wyville Thomson, collected from the general area of the Rockall Trough from 1973 to 1983, were examined; 47% contained material. This varied in nature and in volume between individuals, but overall, the diets of the 6 species showed quite high similarity. The few differences evident are interpreted as reflecting different lifestyles. Except for Ophiocten gracilis, the probably motile, epifaunal species (Ophiura irrorata, O. ljungmani and Ophiomusium lymani) showed the greatest variety in items, and the lowest similarity with the remaining species; they are probably unselective omnivores, eating whatever small prey or organic detritus they are able to find. Ophiacantha bidentata and Ophiactis abyssicola showed both the least dietary variety, and the highest similarity to each other. This probably reflects their more sedentary lifestyle, feeding on current-borne particles and small prey from “perches” on sessile fauna. The stomach contents of Ophiocten gracilis consisted mainly of amorphous organic material and mineral particles, indicating a microphagous feeding mechanism. With O. gracilis, the proportion of stomachs with food decreased markedly in summer, this coinciding with seasonal gonadal development when stomachs frequently appeared ruptured or compressed. The possible importance of seasonal sedimentation of fast-sinking particulates from the surface is discussed in relation to finds of flocculent organic material, often containing diatom frustules, in stomachs of all 6 species. Some doubts remain as to whether our results are biased by specimens egesting stomach contents on capture. As in shallow water, deep-sea ophiuroids seem to be trophic generalists lacking in dietary specialization. Brief notes on parasites encountered are also given.