Small-scale spatial and seasonal differences in the distribution of beach arthropods on the northwestern Tunisian coast. Are species evenly distributed along the shore?

Spatial distributions of sand beach arthropods were studied at Zouara (Nefza), a beach dune system along the northwestern coast of Tunisia. Two transects with pitfall traps, perpendicular to the shoreline and placed from sea towards land, captured spontaneously crawling arthropods. The pitfalls were used to obtain data on horizontal zonations for 2 days during different seasons (April, October). In April, in order to assess local, long-shore distributions of surface-active beach invertebrates, ten transects with pitfall traps were placed every 100 m, for 2 days. Core samples were also taken in correspondence with each of the ten transects to obtain the distribution of burrowing individuals. Sand samples were taken for successive laboratory analyses (salinity, grain size and organic contents). Local climatic conditions were also recorded. One isopod (Tylos europaeus, subdivided into three different size classes), two amphipods (Talitrus saltator, Talorchestia brito and indistinguishable juveniles) and four coleopteran species (Eurynebria complanata, Scarites laevigatus, Phytosus nigriventris, Phaleria acuminata) were considered. Mean zonations perpendicular to the shoreline showed that crustaceans were found closer to the sea-line limits than coleopterans and that between seasons some species changed their mean zonation more than others. Local, long-shore distributions of active and burrowing individuals showed that most species were unevenly distributed along the shore. These discontinuous distributions differed according to the species and age class and were mainly related to specific microclimatic conditions and to nutrition.     

Multiple regression analysis of the sources of variation in orientation of two sympatric sandhoppers, <Emphasis Type="Italic">Talitrus saltator</Emphasis> and <Emphasis Type="Italic">Talorchestia brito</Emphasis>, from an exposed Mediterranean beach

Two sympatric species from an exposed sandy beach in north-western Tunisia, the sandhoppers Talitrus saltator (Montagu) and Talorchestia brito Stebbing, were compared as regards orientation under natural conditions. Sets of experiments were carried out on the beach using two experimental arenas, one permitting the view of both the sky and landscape and the other of the sky only. Replicates were made at different hours of the day, in different days and period of the year and with various climatic conditions. Multiple regression models were fitted to the angular data obtained. These models allow an estimate of the trends of variation depending on a combination of factors. The simultaneous effects of environmental factors influencing orientation were quantified for each species. The sun was confirmed as the major orientation cue in these populations, but the concentration of directional choices was enhanced by the landscape vision. The climatic factors affected orientation in different ways depending on the time of day and year. When the two species were compared, Talitrus saltator showed a higher flexibility of response than Talorchestia brito under the same environmental conditions, supporting the hypothesis of a higher level of terrestrialization of the former species.     

Environmental factors influencing the surface activity and zonation of Tylos europaeus (Crustacea: Oniscidea) on a Tyrrhenian sandy beach

The present study considers a population of Tylos europaeus Arcangeli, 1938 living on a Tyrrhenian sandy beach (Burano, GR, Italy). Monthly surveys were carried out between April 1986 and March 1987 with directional pitfall traps so as to intercept the isopods moving in four directions on the beach surface. In addition a bimonthly study was performed from March 1991 to January 1992 using two methods of capture: pitfall traps joined by 10-cm high strips of fibreglas for surface-active individuals and sieving for those burrowed in the sand. Capture frequencies allowed analysis of annual abundance, daily activity and zonation of juveniles and male and female adults. Variations of these spatio-temporal data were correlated by means of multiple regression with many environmental parameters: temperature and relative humidity of the air and sand, evaporation, wind direction and speed, global radiation, atmospheric pressure, rainfall, the sand salinity and granulometric parameters. T. europaeus was found to be mainly active in summer and autumn and during the night, and was zoned along the eulittoral. The surface activity was influenced by almost all the environmental factors when they were limiting but especially by the relative humidity of the air. The mean zonation of active specimens, however, varied hourly according to the sand temperature. On the other hand, the zonation of the buried individuals depended on the mean grain size, which involves many other parameters, such as moisture and oxygen contents.     

The use of slope and visual information in sandhoppers: innateness and plasticity

Mediterranean sandhoppers of the species Talitrus saltator (Montagu) from the Tyrrhenian population of Burano (Grosseto, Italy) were tested by rearing under different conditions in order to enquire into the innateness and modifiability of orientation with respect to substrate slope and landscape features. Tests were conducted from November 1990 to November 1991. The results were as follows: the sandhoppers responsed to slope only by orienting themselves downslope on dry substrate and upward on a wet one. These responses appeared to be innate but modifiable by experience. Visual dishomogeneity (half horizon covered with a black strip), with a horizontal substrate, elicited an innate orientation towards the black shape, again modifiable by experience. When a black strip was positioned around the upper half of the inclined arena, the sandhoppers responded differently, dependent upon their prior experience: wild individuals responded to the visual stimulus, amphipods reared in optical homogeneity did not make a choice and those reared in conditions identical to those of the test arena responded to slope by orienting downwards and away from the balck strip. With a black strip around the lower half of the inclined arena and with a dry substrate identical responses were shown by wild-caught and laboratory-reared sandhoppers, i.e., downslope and towards the black strip.     

Biomonitoring of polychlorinated byphenyls contamination in the supralittoral environment using the sandhopper Talitrus saltator (Montagu)

We evaluated the use of Talitrus saltator as biomonitor of polychlorinated byphenyls (PCBs) contamination of the supralittoral zone of Mediterranean sandy shores, an area not yet investigated about the presence of these pollutants. Twenty of 22 PCB congeners analysed were detected and quantified in sand and sandhoppers' samples collected in 6 different sites along the Tyrrhenian coast of Central Italy. Among the congeners analysed, PCB 153 and PCB 138 were the most abundant both in amphipods and sand samples and, regarding the congener pattern analysis, hexa-, hepta- and penta-PCBs were the most abundant congeners in sandhoppers. Though hydrophobic compounds, such as PCBs, bind strongly to sand and sediments, PCB concentrations were higher in T. saltator than in the sand revealing a good ability of this species to accumulate this class of contaminants. The concentration of each PCB congener recorded in tissues of sandhoppers showed significant differences among sampling sites revealing a good ability of these species to accumulate PCBs at different concentrations depending on the contamination levels of the sampling site. Therefore, our results suggest the possible utilisation of T. saltator as a biomonitor of PCB contamination of the supralittoral zone of Mediterranean sandy shores.     

Reproductive cycle of two commercial species of sea cucumber (Echinodermata: Holothuroidea) from Caribbean Panama

The reproductive status of the holothuroid species Isostichopus badionotus (Selenka, 1867) and Holothuria mexicana (Ludwig, 1875) was studied over 16 months in Bocas del Toro (Panama), from November 1999 to February 2001. Sexual reproduction was evaluated by the gonad index method, and by histology of gonad development. In addition, population structure was assessed based on sex ratio, minimum reproductive size, and length and weight distributions of males and females. The sex ratio in both species was 1:1, with a unimodal population distribution composed mainly of mature individuals. The minimum reproductive length and weight were 13-20 cm and 150 g, respectively, for both species, although reproductive individuals 10 cm in length were also found. A consistently higher gonad index was observed in H. mexicana, due to a high proportion of mature females and males and high gonad indices in most monthly samples. Gametogenesis and spawning patterns seemed to occur throughout the year, with periods of enhanced activity. Two periods of maximum reproductive activity were tentatively identified: July-November for I. badionotus and February-July for H. mexicana, but neither species had a single, sharply defined annual spawning event. Further work on these exploited holothuroids should examine the relationships between reproduction and environmental factors and between reproductive status and recruitment.     

A histological study of reproduction in the serpulids<Emphasis Type="Italic"> Pomatoceros triqueter</Emphasis> and<Emphasis Type="Italic"> Pomatoceros lamarckii</Emphasis> (Annelida: Polychaeta)

The reproductive cycles of the serpulid polychaetes Pomatoceros lamarckii (Quatrefages, 1865, as Vermilia) and P. triqueter (Linnaeus, 1767, as Serpula triquetra) were studied by histological examination during 1999 and 2000 at Bantry Bay, south-west Ireland. Gametogenesis, sex ratios and hermaphroditism were also investigated. The male/female ratio in P. lamarckii was approximately equal, but in P. triqueter it deviated significantly in favour of females. There was no significant variation in the sex ratios throughout the year in either species. Results confirm that the species are protandric hermaphrodites. In P. lamarckii, but not in P. triqueter, sexual dimorphism was exhibited in body size, the females being larger than the males of the same species. In both species, the proportion of females increased with increasing body size. Simultaneous hermaphroditism was recorded in P. lamarckii, but not in P. triqueter. There was no clearly defined annual gametogenic cycle in either species. Both species appear to have an extended reproductive season, with numerous small-scale peaks in reproductive maturity that can vary annually. Spawning was broadly synchronous between sexes. Visual observation alone (i.e. without histology) was insufficient to accurately assess reproductive condition in individuals not in, or close to, a ripe state.     

Feeding biology and carbon budget of the sediment-burrowing heart urchin Brissopsis lyrifera (Echinoidea: Spatangoida)

This study of the burrowing heart urchin Brissopsis lyrifera includes measurements on feeding and food selection, ingestion rate, absorption efficiency, ventilation and respiration. B. lyrifera regulated feeding depth, ingestion rate and absorption efficiency in relation to food source. When food was added to the top layer of sediment, B. lyrifera burrowed closer to the surface and ingested mainly enriched surface material, whereas it burrowed deeper and ingested deep-seated sediment in the controls. In non-enriched sediment, the feeding rate was 0.04 g sediment DW h^-1 ind.^-1, and, in macro- and microalga-enriched sediment the feeding rate was 0.06 and 0.08 g sediment DW h^-1 ind.^-1, respectively. Absorption efficiency of TOC was about 43% in non-enriched sediment and in microalga-enriched sediment, but was significantly lower (34%) in macroalga-enriched sediment. Absorption efficiency of N varied between 48% and 55%, and was independent of food source. B. lyrifera feed selectively, enriching the gut TOC content about 2-fold and N content about 2.5-fold. The C/N ratio was therefore lower within the gut compared to the surrounding sediment, while the faecal C/N ratio was almost equal to the surrounding sediment. The faeces were, however, slightly richer in TOC and N compared to the surrounding sediment. For 3-5 cm long B. lyrifera, water current rate varied between 4 and 24 ml water h^-1, with a mean of 11 ml h^-1. Mean respiration rate was 205 µl O₂ h^-1 ind.^-1. The water current rate was not sufficient for B. lyrifera to sustain itself by filter feeding only. However, organic-rich particles from the surface are suggested to be an important contribution to the diet. A carbon budget was calculated for B. lyrifera from measured values of consumption, absorption efficiency and respiration, in order to estimate annual production of B. lyrifera. Compared to literature values, growth was overestimated about tenfold in the budget. A large proportion of the absorbed carbon was suggested to leave the animal as dissolved carbon, through mucus production or through anaerobic pathways, either by the heart urchin or by micro-organisms in the gut.     

Astronomical, landscape and climatic factors influencing oriented movements of Talitrus saltator in nature

 The present study was conducted on a Mediterranean beach (Burano, southern Tuscany, Italy) to examine the timing, orientation, and motivating and directing factors of the spontaneous movements of the sandhopper Talitrus saltator (Montagu, 1808). In April 1994, October 1994 and June 1995, during different moon phases, traps were positioned in the eulittoral zone, which intercepted sandhoppers when walking on the sand surface. At the same time environmental parameters were registered. Contemporaneously, orientation tests were carried out on active individuals using two arenas, one of which permitted a view of both sky and landscape and one which prohibited the landscape view. The results show landward migration after sunset, for juveniles later than for adults, nonoriented activity for 2 or 3 h after midnight and seawards zonal recovery before and after sunrise. The main microclimatic factor modulating activity was sand temperature, while the nonoriented activity between the two migratory movements seems to be endogenously determined. Both sky and landscape cues are used by sandhoppers for orienting their course, but the landscape view is sufficient and necessary on new moon nights. These results contribute to solution of the controversy on the mechanisms actually motivating sandhoppers in nature. Received: 24 September 1996 / Accepted: 25 October 1996     

Interaction between solar orientation and landscape visibility in Talitrus saltator (Crustacea: Amphipoda)

A series of experiments on the littoral amphipod Talitrus saltator (Montagu) was carried out between April and September, 1978–1981, both under natural conditions and inland with artificial landscapes of different heights on the horizon, in order to assess the visual importance of the landscape in zonal orientation in populations from the Mediterranean coast, and to determine interactions between solar orientation and orientation based on the landscape. Inland, orientation in controls (permitting only vision of the sky and the sun) was compared to that of the experimental individuals, who could see a simulated landscape positioned landwards to them and seawards. In nature, sandhoppers released in a level arena with the landscape screened from view were compared with others released in an unscreened arena and on the sand in absolutely natural conditions. Situations where solar orientation contradicted local cues were produced both by releasing the sandhoppers on a shore diversely orientated to their own, and by shifting their internal clock by nine hours. Results show that T. saltator uses the landscape as a cue in its orientation towards the sea, in conjunction with solar orientation: the latter being the principal factor involved even when the sandhoppers are separated from the local optical factor in the sky (in trials at a distance from the sea). In fact, with the artificial landscape set seawards, none of the populations we studied showed any orientation based on the landscape comparable in accuracy to orientation based exclusively on the sun. From the experiments carried out in natural conditions, it is possible to deduce that conflicting conditions gave rise to two types of results: deviation from the mean direction and an increase in dispersion, up to total dispersion when the local factors were in total contrast to solar orientation (clock-shifted sandhoppers released on the sand).     

Surface activity,zonation and migrations of Talitrus saltator on a Mediterranean beach

The present study considers a population of Talitrus saltator (Montagu, 1808) living along a Mediterranean beach (Burano, Grosseto, Italy). From April 1986 to March 1987 the annual and daily patterns of amphipod activity were analysed together with zonation and the direction and extent of movements. An annual bimodal activity pattern was established with a first peak of activity in late spring and another one in autumn. Daily activity was predominantly nocturanl with two peaks, one of which could be suppressed or depressed in accordance with particular climatic conditions. The species studied was generally limited to the eu- and supralittoral part of the beach, but the extent of zonation could vary according to climate. Differences between juveniles and adults appeared both for activity patterns and for zonation. Movements were predominantly (ca. 85%) along the sea/land axis with a landward migration always followed by seaward one. Under conditions of high relative humidity and rain, the migrations towards land could extend to the dune. Movements parallel to the shoreline were demonstrated in a separate experiment (in June and October 1987 and June 1988) using the capture-mark-recapture method. The maximum distance covered registered in one night was 200 m.     

Spatial and temporal strategies in the surface activity of some sandy beach arthropods living along the French Atlantic Coast

During a field study on the eco-ethology of sandy beach arthropods conducted along the French Atlantic Coast it emerged thatTylos europaeus, Phaleria cadaverina andLabidura riparia were among the most abundant species that lived in the eulittoral. Tetradirectional pitfall traps, placed from the base of the dune to the tidal limits, were used. The traps intercepted surface-active arthropods, and the number of traps varied according to the levels of the tides. Data were recorded hourly during two periods corresponding to a spring tide and the following neap tide. At the saine time the most important environmental parameters were registered each hour. The results indicate that all three species show prevalently nocturnal surface activities. ForT. europaeus andPh. cadaverina an influence of the synodic and tidal phases was found on the zonation of the surface activity and on the extent of movement. The latter was due to the greater or smaller distances that the isopods and tenebrionids had to move from the diurnal burrowing sites to the foraging zones. ForL. riparia no variations in the total mean zonations were found between the two synodic phases, and only at neap tide were seaward, followed by landward movements found. Multiple regression analysis showed that the environmental parameters registered in this period were never limiting, as threshold values in this case probably were not reached.     

Biological patterns and near-bottom population characteristics of two pasiphaeid decapod crustacean species, Pasiphaea sivado and P. multidentata, in the north-western Mediterranean Sea

Biological aspects of the only two epi-/mesopelagic pasiphaeid species distributed along the continental marginof the north-western Mediterranean Sea were studied over the period 1991-1994. The annual reproductive cycle, reproductive output, sex and size distributions by depth (near-bottom distribution), and growth were analysed using 4156 specimens of Pasiphaea sivado (Risso, 1816) and 5491 specimens of P. multidentata Esmark, 1866. The reproductive cycle in the shallower-dwelling species, P. sivado, was nearly continuous, in contrast to the reproductive cycle in P. multidentata, which was markedly seasonal in nature. Both species exhibited similar size and population structure patterns with depth, with individual size increasing along the bottom depth gradient. The shallower-dwelling species had a significantly higher reproductive output level, in terms of the relative number of eggs, than the deeper-dwelling species. Biological trends for each of the species have been compared. The comparative results for these two congeneric species revealed that the biological parameters analysed were closely linked to their population structure. The slight differences in the depth range inhabited by each species have been considered the main environmental factor affecting the life histories of these two species. The data presented supported the hypothesis that depth exerted an important influence on the biological patterns of deep-water species, with the seasonality of reproductive processes increasing in deeper-dwelling species.     

Estimation of age in the sex-changing, coral-inhabiting snail Coralliophila violacea from the growth striae on opercula and a mark–recapture experiment

Change of sex in the coral-inhabiting snail Coralliophila violacea (Lamarck) may occur in a wide range of sizes in the field. One proposed explanation for this is that the snails change their sex at a certain age and that individuals have different growth rates caused by microhabitat differences. In this study, we attempt to establish a method to determine the age of this snail and age at sex change. The growth striae on the operculum were studied and compared to the age estimated by the Gompertz growth function based on growth data obtained from mark-recapture experiments in southern Taiwan. There is a significant correlation between the number of striae on the operculum and the age estimated from the Gompertz growth function, and the relationship is 1:1. These results suggest that the number of striae on the operculum can be used as an age index, with each stria representing 1 year of age. The age of sex change of this snail, according to our estimates by both stria number and aperture-length inferences, occurs between 4 and 6 years old. Growth rates of the snails are negatively correlated to size. Furthermore, individuals undergoing sex-change grow faster than males and females.     

Field evidence of the impact of two acanthocephalan parasites on the mortality of three species of New Zealand shore crabs (Brachyura)

The effects of the acanthocephalan parasites Profilicollis antarcticus and P. novaezelandensis on the fecundity and mortality of three species of shore crab (Macrophthalmus hirtipes, Hemigrapsus edwardsi and H. crenulatus) are examined. The number of eggs produced by female crabs was strongly correlated (all P<0.05) with carapace width; parasite load was not a significant determinant of female fecundity. Mortality was inferred from reduction in the mean number of parasites per crab in the largest crab size-classes, indicating that heavily infected individuals are removed from the population. Mortality attributable to the parasites was observed for all three species of crabs, although the effect of parasites varied in both time and space; significant curvilinear regressions between parasite load and crab size-classes were not found in all samples. Crab mortality appears to be influenced by more than the pathological influences exerted by the parasites. Parasite-induced behavioural alterations may cause crab hosts to be more susceptible to predation by definitive hosts. We support this suggestion with three lines of evidence: the lack of parasite effects on fecundity, the weakening of the parasite effect on mortality during the time of year when birds are absent, and previous indications of parasite-mediated alterations in crab burrowing behaviour.     

Activity patterns of Kemp's ridley turtles, <Emphasis Type="Italic">Lepidochelys kempii</Emphasis>, in the coastal waters of the Cedar Keys,Florida

Radio and sonic telemetry were used to investigate the tidal orientation, rate of movement (ROM), and surfacing behavior of nine Kemp's ridley turtles, Lepidochelys kempii, tracked east of the Cedar Keys, Florida. The mean of mean turtle bearings on incoming (48ᇅ °) and falling (232ᆽ °) tides was significantly oriented to the mean directions of tidal flow (37Nj °, P<0.0025, and 234Nj °, P<0.005, respectively). Turtles had a mean ROM of 0.44ǂ.33 km/h (range: 0.004-1.758 km/h), a mean surface duration of 18ᆣ s (range: 1-88 s), and a mean submergence duration of 8.4Lj.4 min (range: 0.2-60.0 min). ROM was negatively correlated with surface and submergence durations and positively correlated with the number of surfacings. Furthermore, ROMs were higher and surface and submergence durations were shorter during the day. Daily activities of turtles were attributed to food acquisition and bioenergetics.     

Gonad morphology, sexual development, and colony composition in the obligate coral-dwelling damselfish Dascyllus aruanus

Gonad morphology and colony composition support the existing supposition that the obligate coral-dwelling damselfish Dascyllus aruanus has a protogynous hermaphroditic sexual pattern. Adults had either an active ovary containing vitellogenic oocytes, an ovotestis, or a spermiated testis and were classified as adult female, hermaphrodite, or adult male, respectively. Among individuals having male function, the testis (or testis portion of the ovotestis) takes the form of an unrestricted spermatogonial lobular testis. Among hermaphrodites having an ovotestis, a small proportion of individuals had a gonad in which both the ovarian and testicular portions were inactive (inactive hermaphrodites), whereas the majority had a predominantly testicular ovotestis that contained spermatozoa (male-active hermaphrodites). The size range of individuals within gonadal classes indicates that all D. aruanus first develop an ovariform gonad. Some individuals then undergo ovarian maturation to become adult females while others develop testicular tissue to form an ovotestis and become male-active hermaphrodites. Subsequently, progressive loss of ovarian tissue results in the development of a secondary testis from an ovotestis with the retention of a residual, afunctional lumen among adult males. The wide size range of individuals having an ovotestis suggests that some hermaphrodites function as adult females before developing testicular tissue while other individuals do not pass through an adult female stage. If this is the case, D. aruanus exhibits a diandric protogynous hermaphroditic sexual pattern. The apparent prolonged retention of an ovotestis with both healthy oocytes and an ovarian-type lumen in a spermiated ovotestis, as well as a functional sex ratio of 1:1 for adult females:adult males plus male-active hermaphrodites also raises the possibility that D. aruanus may be capable of bidirectional sex change during the hermaphroditic stage. Such a capability would be highly adaptive for a species having limited mobility and unpredictable recruitment of new colony members resulting in unpredictable mating opportunities.     

Origin of male-biased sex allocation in orphaned colonies of the termite, <Emphasis Type="Italic">Coptotermes lacteus</Emphasis>

In eusocial insects, sex allocation often constitutes a ground for intracolonial conflicts. This occurrence provides ideal opportunities to test kin-selection theory. A vast literature on this topic is available for social Hymenoptera, but the same field remains almost untouched in termites. A preeminent case is that of some species of Coptotermes, where the sex-allocation ratio in nymphs shifts from near equity to all-male when the primary reproductives are replaced by neotenics. To shed light on the developmental origin of this shift, we compared the sex ratio of the various castes and instars in primary- and neotenic-headed mature colonies of Coptotermes lacteus. The male-biased sex allocation in the latter type of colony results from two concurrent events: first, the sex ratio of the youngest instars (larvae) is male-biased by a 3:1 ratio; and second, all female larvae become workers, while a large fraction of the male larvae proceed to the nymphal and alate stages. Colony-founding experiments showed that inbreeding by itself cannot account for the male bias at hatching. We suggest that both genetic factors, due to the reproductive behaviour of neotenics, and environmental factors (colony condition and resource availability) may influence this process. Their exact nature and respective impact have not yet been clarified.     

Egg production in the arrow squid Nototodarus gouldi (Cephalopoda: Ommastrephidae), fast and furious or slow and steady?

The process of reproductive maturation and egg release was examined in the temperate shelf squid Nototodarus gouldi. The energy allocation between somatic and reproductive growth from juvenile to mature adult was investigated throughout the life span to determine the underlying energetic strategy adopted by individuals. The relative weight of the mantle, fin and digestive gland remained unchanged during ovarian development, with no significant correlations found between the mantle length (ML)-gonad residuals and the ML-mantle (r=0.01, P>0.05), ML-fin (r=0.07, P>0.05) and ML-digestive gland (r=0.07, P>0.05) residuals. This suggested that energy was not being diverted away from somatic growth during sexual development, and consequently neither muscle nor digestive gland was being utilised as an energy store. Since squid in all maturity stages contained some food in their stomachs (e.g. 66.7% of mature animals), it is likely that the cost of maturation in this species is largely being met by food intake. The energy investment in reproductive tissues was relatively low (mean gonado-somatic index for mature individuals was 9.29% - 0.40%), indicating that only small amounts of energy were being allocated to reproduction at anyone point in time, which is characteristic of a multiple-spawning strategy. Furthermore, oviduct weight was not correlated with body size (r=0.256, P>0.05), suggesting that eggs are not stored for a single release. In all except one individual, ovary weight was consistently heavier than oviduct weight, suggesting that the ovary is not being depleted of oocytes as mature ova move into the oviducts. Additionally, the ovaries of mature females contained a range of oocyte sizes with discrete peaks, indicating a continued production and development of oocyte cohorts. The presence of some individuals with stretched empty oviducts is further evidence that the reproductive strategy of N. gouldi is slow and steady, with eggs possibly being released in discrete batches over a period of time.     

Burrowing behaviour and sediment reworking in the heart urchin Brissopsis lyrifera Forbes (Spatangoida)

Sediment reworking due to burrowing and feeding was studied in the spatangoid Brissopsis lyrifera, at two different temperatures (7°C and 13°C). Spine activity and burrowing behaviour were recorded with a real-time video camera. Reworked sediment volume was calculated from tracks produced by the heart urchin. Ingestion rates were measured by feeding the heart urchins with luminophores. Temperature had a significant effect on the bioturbation activity of B. lyrifera. At 13°C reworked sediment volume due to burrowing was 22 ml sediment h^-1 and at 7°C 14 ml sediment h^-1. The ingested amount of sediment was 0.08 and 0.02 g dry sediment h^-1 in 13 and 7°C, respectively. Reworked sediment volume due to burrowing was 60-150 times higher than the volume ingested. The large reworked volume is a consequence of B. lyrifera moving with a rocking motion through the sediment. The spines were continuously transporting sediment around the test with 5-min metachronal wave cycles.