Depth regulation of larval marine decapod crustaceans: test of an hypothesis

This study tested the hypothesis that the dimensions and symmetry of the depth regulatory window of crustacean larvae are controlled by the level of light adaptation. Responses of first and last zoeal stages of the crab Rhithropanopeus harrisii (Gould) to different rates of pressure change were analyzed with a video system. Crabs were collected from the Neuse River estuary (North Carolina, USA) from May to September 1988. Responses were measured when larvae were adapted to light having an angular light distribution similar to that underwater at intensities ranging from one log unit above the lower phototaxis threshold to four log units higher. For both zoeal stages in darkness and at 10^-6 W m^-2, the distance larvae descend before responding to a pressure increase was much shorter than the distance they would ascend before responding to a pressure decrease. When adapted to a light level of 10^-4 W m^-2 both zoeal stages descended and ascended approximately equal distances before responding to an increase or decrease in pressure, respectively. Finally at the highest test light intensity (10^-2 W m^-2), the ascent distance was much shorter than the descent distance. These results support the hypothesis. The depth regulatory window dimensions predict an ascent in the water column upon adaptation to low light intensities and descent at high light levels. Thus Sulkin's negative feedback model provides the general mechanism of depth regulation. The effects of light adaptation on the limits of the depth regulatory window provide an additional component that negates the requirement for depth regulation at an absolute depth. The composite model can be termed the light-dependent negative feedback model of depth regulation.     

Comparative study of crustacean larval photoresponses

Ovigerous females of four brachyuran (Cancer gracilis, Lophopanopeus bellus bellus, Hemigrapsus oregonensis and Scyra acutiforns) and two anomuran (Pagurus beringanus and P. granosimanus) species, which live as adults in coastal areas, were collected near Friday Harbor, Washington, USA, in 1985, and spectral sensitivity and phototactic pattern of their larvae were measured. Responses were compared with previous measurements on estuarine species to determine whether responsiveness varies with adult habitat. Estuarine and coastal species have similar photoresponses. Spectral sensitivity of the test brachyran species had two maxima, one near 400 to 420 nm and another around 500 nm. The anomuran species were similar, but had an additional peak in the region of 580 to 620 nm. This sensitivity is adapted to daytime light conditions in the adult environment, and available spectra at the time of larval movement during diel vertical migration. Upon light adaptation and stimulation with a narrow light field, all six species showed positive phototaxis to high light intensities and a pronounced negative response to low intensities. Only the negative response will occur in natural underwater light conditions, and it is part of a predator-avoidance shadow-response which operates in areas of higher light intensity. The same phototactic pattern is observed in all species when darkadapted except H. oregonensis. If nocturnal vertical migration occurs, this negative response may be responsible for the descent at sunrise and depth maintenance during the day.     

Occurrence of a shadow response among brachyuran larvae

A closed-circuit television system was used to study phototaxis and behavioral responses upon sudden decreases in light intensity in light-adapted Stage I zoeae from 7 species of Brachyura (Panopeus herbstii, Menippe mercenaria, Uca pugilator, Callinectes sapidus, Sesarma cinereum, Pinnotheres maculatum, and Libinia emarginata). All species except one show positive phototaxis to high intensities of 500 nm light and negative phototaxis to low intensities. Upon a sudden light intensity decrease, larvae show a shadow response, which depends upon the initial and final intensities. If the initial intensity is sufficient to induce positive phototaxis, and the light is extinguished, the larvae of 6 species stop swimming and passively sink (sinking response). Reducing the intensity to an absolute level that causes negative phototaxis, induces a sinking response followed by negative phototaxis. For intensity reductions that occur in the intensity range above that for negative phototaxis, a sinking responseis observed. The intensity decrease that initiates the sinking response is independent of initial stimulus intensity and duration, is consistent between species, and is equivalent to a decrease by 0.5 OD (optical density) units. The sinking response is greatest at intensity decreases of at least 1.0 to 1.1 OD units. It is concluded that the shadow response is common in brachyuran larvae inhabiting coastal-estuarine areas.     

Behavioral responses of a larval crustacean to hydrostatic pressure: Rhithropanopeus harrisii (Brachyura: Xanthidae)

Responses of the four zoeal stages of the crab Rhithropanopeus harrisii (Gould) to step and continuous changes in hydrostatic pressure were analyzed with a video system. Crabs were collected from the Neuse River estuary (North Carolina, USA) from June to August, 1987. The lower thresholds for step increases and decreases in pressure were 3 and 8 to 10 mbar, respectively. There was little change in sensitivity with zoeal development. Tests of larval responses in a light field that simulated the underwater angular light distribution indicated positive phototaxis does not occur upon pressure changes. In darkness, rates of pressure increase at and above 0.175 mbar s^-1 induced high barokinesis and negative geotaxis in all but Stage IV zoeae, which had a threshold of 1.19 mbar s^-1. Since larval sinking and descent swimming speeds exceed these threshold rates, larvae can move rapidly enough to produce suprathreshold changes in pressure which evoke behavioral responses. Slow rates of pressure decrease induced passive sinking while rapid rates caused an active ascent. This ascent response upon a pressure decrease is unreported among crustaceans, and is hypothesized to function for avoidance of feeding and respiratory currents of benthic invertebrates. The descent response occurs in all zoeal stages, except IV, at rates of pressure decrease (0.4 to 0.71 mbar s^-1) that are within the range of ascent swimming speeds. These results support Sulkin's negative feedback depth regulation model. The absolute distances moved before corrective vertical responses to threshold rates of pressure change are initated delimit the depth regulatory window. In darkness, the asymmetry of the window would lead to an ascent. It is hypothesized that light is an additional component in depth regulation, and that the limits and symmetry of the depth regulatory window may be controlled by the level of light adaptation.     

Photosensitivity of the calanoid copepod Acartia tonsa

The light intensity and spectral sensitivities of the calanoid copepod Acartia tonsa Dana were determined by measuring phototactic responses. Adult females displayed only positive phototaxis. The dark-adapted copepod, which possesses a single naupliar eye, perceived light at intensities as low as 2.8x10¹¹ photons m^-2 s^-1. The action spectrum for positive phototaxis had no clear maxima but rather showed a broad range of greatest sensitivity from 453 to 620 nm. This sensitivity encompassed those wavelengths that are maximally available at the depth where the copepod is found during the day. This spectral overlap, coupled with the finding that the copepod requires light cues for nocturnal vertical miration, suggests that broad spectral sensitivity is an adaptive mechanism to maximize light intensity sensitivity during migration.     

Bioluminescence in the Monterey Submarine Canyon: image analysis of video recordings from a midwater submersible

Video images of bioluminescence were recorded in situ during a 1985 study of the midwater environment of the Monterey Canyon, using a single-person, untethered submersible. Gelatinous organisms were responsible for the most brilliant bioluminescent displays, often exhibiting elaborate kinetics in response to mechanical stimulation. Images of bioluminescent displays recorded from identified organisms are shown and display patterns are described. All bioluminescence emission spectra from captured specimens were blue, with peak emissions between 460 and 494 nm. Image-analysis of recordings of mechanically stimulated bioluminescence revealed source densities between 43 and 175 m^-3 and intensities between 2.5 and 37.3 W sr^-1 m^-3. The predominant display type at all depths studied (between 100 and 560 m) was luminous secretions. Despite high intensities of mechanically stimulated bioluminescence, no spontaneous light production was recorded in the absence of mechanical stimulation.     

Patterns of growth and triacylglycerol content in snow crab Chionoecetes opilio (Brachyura: Majidae) zoeal stages reared in the laboratory

Snow carb Chionoecetes opilio zoea I and zoea II larvae, hatched from females in a controlled mating experiment, were reared in the laboratory at 10.1 °C and 28.0 salinity, to resolve the patterns of growth (dry weight [DW]) and change in energy reserves (triacylglycerols [TAG]) within a given moult cycle. The patterns of growth and change in TAG reserves were similar in each zoeal stage. Following hatching or a moult, the zoeae entered a phase of rapid size increase, i.e. high daily growth rates (5.5 to 12.8% DWd^-1), for 1/3 to 2/5 of the duration of the moult cycle. During the same period, the zoeae accumulated TAG reserves until a maximum (TAG DW^-1) was reached at the end of the phase of rapid growth. The period of high growth rates and of TAG accumulation is interpreted as the required time for the zoeae to reach a point in development [i.e. point of reserves saturation (PRS); Anger and Dawirs (1981)] where sufficient growth and energy reserves allow moulting to the next stage. Following the phase of rapid growth and TAG accumulation, the zoeae entered a phase of low daily growth rates (0 to 1% DWd^-1) during which the TAG reserves decreased to a minimum at the end of the phase. Prior to, and during the moult to zoea II, a phase of negative growth was observed in the zoea I larvae. We conclude that measurement of zoeal size and TAG content, along with morphometric criteria (e.g. epidermal retraction), can be used to assess growth and nutritional condition of C. opilio zoeal stages from the sea.     

Biochemical changes during larval development of the xanthid crab Rhithropanopeus harrisii. I. Protein,total lipid,alkaline phosphatase,and glutamic oxaloacetic transaminase

Protein, total lipid, and the activities of the enzymes alkaline phosphatase (AP) and glutamic oxaloacetic transaminase (GOT) were measured daily during larval development of the xanthid crab Rhithropanopeus harrisii (Gould). Percent increase in fresh weight was greatest on the first day after each molt. Protein and total lipid per larva increased during the second, third, and fourth zoeal stages, with the most marked increase during the third. When calculated as percent fresh weight, marked accumulation of both protein and total lipid occurred during the third zoeal stage. GOT activity remained stable during the third zoeal stage, but increased significantly during the fourth zoeal stage. AP showed a 4-day cycle of activity apparently related to the molt cycle. It is suggested that increased protein and total lipid content during the third zoeal stage indicate nutritional requirements prior to metamorphosis. Biochemical change may be a more relevant indicator of growth during larval development than either size increase or molt frequency when assessing the value of various diets in the culture of crabs.     

Facultative lecithotrophy during larval development of the burrowing shrimp Callianassa tyrrhena (Decapoda: Callianassidae)

Survival, developmental and consumption rate (Artemia nauplii ingested per day) as well as predation efficiency (ingested per available Artemia nauplii) were studied during the larval development of the shallow-water burrowing thalassinid Callianassa tyrrhena (Petagna, 1792), which exhibits an abbreviated type of development with only two zoeal stages and a megalopa. The larvae, hatched from berried females from S. Euboikos Bay (Aegean Sea, Greece), were reared at 10 temperature–food density combinations (19 and 24 °C; 0, 2, 4, 8 and 16 Artemia nauplii d⁻¹). Enhanced starvation resistance was evident: 92 and 58% of starved zoeas I molted to zoea II, while metamorphosis to megalopa was achieved by 76 and 42% of the hatched zoeas at 19 and 24 °C, respectively. The duration of both zoeal stages was affected by temperature, food density and their interaction. Nevertheless, starvation showed different effects at the two temperatures: compared to the fed shrimp, the starved zoeae exhibited accelerated development at 19 °C (8.4 d) but delayed metamorphosis at 24 °C (5.9 d). On the other hand, both zoeal stages were able to consume food at an increased rate as food density and temperature increased. Predation efficiency also increased with temperature, but never exceeded 0.6. Facultative lecithotrophy, more pronounced during the first zoeal stage of C.tyrrhena, can be regarded as an adaptation of a species whose larvae can respond physiologically to the different temperature–food density combinations encountered in the wide geographical range of their natural habitat. Received: 28 February 1998 / Accepted: 21 October 1998     

Foraging behaviour of larval cod (<Emphasis Type="Italic">Gadus morhua</Emphasis>) at low light intensities

The ability to forage at low light intensities can be of great importance for the survival of fish larvae in a pelagic environment. Three-dimensional silhouette imaging was used to observe larval cod foraging and swimming behaviour at three light intensities (dusk ~1.36 × 10⁻³ W/m², night ~1.38 × 10⁻⁴ W/m² and darkness ~3.67 × 10⁻⁶ W/m²) at 4 different ages from 6 to 53 days post-hatch (dph). At 6 dph, active pursuit of prey was only observed under dusk conditions. Attacks, and frequent orientations, were observed from 26 dph under night conditions. This was consistent with swimming behaviour which suggested that turn angles were the same under dusk and night conditions, but lower in darkness. Cod at 53 dph attacked prey in darkness and turn angles were not different from those under other light conditions. This suggests that larvae are still able to feed at light intensities of 3.67 × 10⁻⁶ W/m². We conclude that larval cod can maintain foraging behaviour under light intensities that correspond to night-time at depths at which they are observed in the field, at least if they encounter high-density patches of prey such as those that they would encounter at thin layers or fronts.     

Effects of the insect growth regulator Dimilin® (TH 6040) on larval development of two estuarine crabs

Effects of Dimilin^® (TH 6040), an insect growth regulator which interferes with the formation of the insect cuticle, were studied on the larval development of Rhithropanopeus harrisii (Gould) and Sesarma reticulatum (Say) (Crustacea: Brachyura). When larvae were exposed to 0.5 (R. harrisii only), 1, 3, 5, 7, and 10 ppb Dimilin from hatching to the first crab stage, survival in both species decreased in relation to increased concentrations of Dimilin. Survival of R. harrisii larvae wa significantly lower at 1 ppb and higher levels compared with control experiments, and in S. reticulatum a significant decrease in survival began at the 3 ppb level. At 10 ppb Dimilin, no larvae survived to the megalopa stage in either of the two species. The results indicate that early stage larvae of R. harrisii are more sensitive to Dimilin than those of S. reticulatum. When R. harrisii larvae were treated with 10 ppb Dimilin during the intermolt period of each of the 4 zoeal stages, nearly all larvae died during molting to the succeeding stage. First zoeal larvae of R. harrisii exposed to 10 ppb Dimilin at various days during the intermolt period were more sensitive to the compound late than early in the period. It is suggested that Dimilin also may interfere with the formation of the cuticle in crab larvae.     

Larval settlement and metamorphosis in the sand crab Emerita talpoida (Crustacea: Decapoda: Anomura)

The effects of various potential cues on the survival and time to metamorphosis of larvae of the sand crab Emerita talpoida (Say) were examined. Zoeal duration, which ranged from 25 to 43 d, was not correlated with subsequent megalopal survival but had a weak, negative correlation with the duration of the megalopal stage. Neither food, sand nor water from containers with adults directly affected megalopal survival or duration. In a second experiment, the presence of sand both increased the probability of survival and shortened the duration of late zoeal stages. This is the first experimental evidence for delayed zoeal metamorphosis in a decapod. Settlement of E. talpoida appears to occur primarily during the zoeal phase, not at the megalopal stage.     

UV- and salinity-induced oxidative effects in the marine diatom <Emphasis Type="Italic">Cylindrotheca closterium</Emphasis> during simulated emersion

The diatom Cylindrotheca closterium was exposed to transient light- and osmotic conditions as occur during its tidal emersion. The objective was to analyze how this simulated emersion contributes to the production of active oxygen species (AOS) and via this, to oxidative cell damage. Light- and salinity conditions were varied in factorial combination: low light (no UVB) or high light (unweighted UVB-dose rates of respectively 0.01; 0.07; 0.24; 1.03 W m⁻²) at normal (30 psu) or high salinity (60 psu). UVB (0.01–0.24 W m⁻²) and high salinity had a significant, negative effect on the photosynthetic efficiencies ΔF/F _m’ (steady-state quantum yield) and F _v/F _m (maximum yield). UVB at 1.03 W m⁻² (15 kJ m⁻² d⁻¹) almost arrested electron transport. At ecologically relevant UVB levels, i.e. below 0.24 W m⁻² (≈3.4 kJ m⁻² d⁻¹) with UVB:PAR<0.4:100 (PAR photosynthetically active radiation) only dynamic photoinhibition was observed (protection via heat dissipation). Non-photochemical quenching was positively correlated with the de-epoxidation of diadinoxanthin (DD) to diatoxanthin (DT). A decreasing ratio DT/(DD+DT) after 4 h of UVB at >0.07 W m⁻² and at 60 psu indicated a reversal of the diatom xanthophyll cycle (diminished photoprotection) which may be caused by an enhanced AOS production. Oxidative stress and -damage to C. closterium cells were assessed applying fluorescent indicator dyes, via confocal microscopy and quantitative image analysis. AOS production rates (cellular DCF fluorescence) were stimulated by UV, and were ~50% higher at 60 psu. AOS production decreased with an increasing pre-exposure (0–4 h) to normal UVB (0.24 W m⁻²), which indicated a stimulation of the antioxidative defence. Non-protein thiols (indicator CMF) and glutathione pools (HPLC-analyzed) decreased with UVB-dose rates (0.01–0.24 W m⁻²), most likely due to AOS-mediated thiol oxidation. Hypersalinity (60 psu) and UVB (0.01–0.24 W m⁻²) caused membrane depolarization (dye DIBAC₄(3)) and phospholipid hydrolysis (phospholipase A₂ dye: bis-BODIPY FL-C₁₁-PC). AOS production may have diminished the membrane polarity, and peroxidized the membrane lipids (HPLC-analyzed malondialdehyde) which enhanced PLA₂ activity. The dyes indicated an increased oxidative (lipid) damage at a 15% inhibition of photosynthesis in this diatom, at UVB levels and salinities that can be expected in situ during its periodic tidal emersion.     

Escape speeds of marine fish larvae during early development and starvation

Response rates to tactile stimulation and subsequent escape speeds were measured using a video-recording system during early development and starvation of fish larvae. The species studied included the yolk-sac larvae of Clyde and Baltic herring (Clupea harengus L.), cod (Gadus morhua L.), flounder (Platichthys flesus L.) and older larvae of Clyde herring. The proportion of larvae responding (response rate) was initially about 20 to 25% in herring and 35 to 40% in cod and flounder using a probe, but about 70 to 80% using the sucking action of a pipette in all species except flounder. Both response rates and escape speeds (mean and maximum) tended to peak 1 to 2 d before the PNR (point-of-no-return, when 50% of larvae are too weak to feed), then decreased slowly during further starvation. An inter-species comparison showed that the highest recorded mean escape speeds (measured over a period of 200 ms) and highest maximum escape speeds (over 20 ms) ranged from 5.7 to 8.6 BL/s (body lengths/s) and 12.1 to 16.1 BL/s, respectively. The larvae made directional responses away from the stimulus only when they developed and reached the feeding stage.     

Energetics of underwater swimming in the great cormorant (Phalacrocorax carbo sinensis)

Resting metabolic rate (RMR), energy requirements and body core temperature were measured during underwater swimming in great cormorants (Phalacrocorax carbo sinensis) at the zoological garden in Neumünster, Germany, using gas respirometry and stomach temperature loggers. We used a 13 m long still water canal equipped with a respiration chamber at each end. Birds swam voluntarily in the canal at a mean speed of 1.51 ms^-1. Power input during underwater swimming averaged 31.4 W kg^-1. Minimal costs of transport of 19.1 J kg^-1 m^-1 were observed at a speed of 1.92 m s^-1. Body core temperature was stable in all birds within the first 60 min spent in the canal. After that, body temperature dropped at a rate of 0.14°C min^-1 until the birds voluntarily left the water. Our data indicate that great cormorants spend 2.7 times more energy than Adélie penguins (Pygoscelis adeliae) during underwater swimming. This can be essentially attributed to their poor insulation, their mode of locomotion underwater and differences in streamlining. RMR on land was related to body mass via VO₂=0.691 M^0.755 (where VO₂ is O₂-consumption in litre h^-1 and M is body mass in kg). In order to quantify the effects of external devices on energy consumption during underwater swimming, we tested a dummy data logger attached to the back of the cormorants as well as a ring on the leg. The ring had no apparent influence on the swimming energetics of the cormorants. In birds equipped with dummy loggers, swimming speed was not significantly influenced, but both power input and costs of transport increased by a mean of 19% for swimming speeds between 1.4 and 1.8 m s^-1.     

Laboratory study of survival and duration of individual zoeal stages as a function of temperature in the brachyuran crab Cancer magister

Larvae were hatched from ovigerous Dungeness crabs, Cancer magister, collected from Puget Sound Basin, Washington, USA, in April, 1986, and the effects of temperature on rates of survival and development were studied for each of the five zoeal stages both in small batch-culture and in individual culture. Culture method had little effect on the results at 10°, 15°, and 20°C. Increased mortality was measured at all stages at 20°C, with 100% mortality occurring during the terminal fifth stage. Fifth stage larvae may also show higher mortality at 15°C than at 10°C. Stage duration varied inversely with temperature at all stages, although differences between 10° and 15°C were greater than between 15° and 20°C. The results indicate that survival and stage duration are independent of the values for the previous and subsequent stages, that variability among larvae in instar duration increases with temperature, and that the terminal fifth zoeal stage is the most sensitive to temperature stress. Duration of a late zoeal instar is not related to its earlier development rate nor can early development rates be used to predict whether individual zoeae will successfully develop to the megalopa. Measurements of megalopa dry weights indicate no differences due either to previous culture temperatures or to total time to the megalopa. Predictive models of larval transport that require estimates of larval duration should account for both changes in temperature response that can affect individual stage duration, and variability among individuals in stage duration that can influence the degree of larval dispersion.     

Effects of the juvenile hormone mimic ZR-515 (Altosid®) on larval development of the mud-crab Rhithropanopeus harrisii in various salinities and cyclic temperatures

Effects of 0.01, 0.1 and 1.0 ppm methoprene (Altosid^®: ZR-515), a juvenile hormone (JH) mimic which shows high activity against some economically important insect pests, especially Diptera, were tested on larvae of the mud-crab Rhithropanopeus harrisii (Gould) (Brachyura: Xanthidae) from hatching to the first crab stage under optimum and stress conditions of a number of salinities and cyclic temperatures. There was a significant reduction in survival of zoeal larvae with increasing concentrations of methoprene in nearly all combinations of salinity and temperature. On the average there was 9% less survival in the 0.01 ppm concentration of methoprene than in the control, and in the 0.1 ppm concentration the survival was further reduced by another 16%. At 1.0 ppm methoprene no larvae survived beyond the first zoeal stage under optimum conditions or under stressful combinations of salinity and temperature. Except at 0.2 ppm in 27.5% S, survival of the megalopa was not significantly reduced in 0.01 or 0.1 ppm methoprene in any salinity or temperature, although the percentage of abnormal megalopa increased under stress conditions. The first zoeal stage was the most sensitive of the larval stages to methoprene as well as to salinity and temperature stress. The duration of zoeal development was significantly lengthened with an increase in concentration of methoprene under nearly all conditions of salinity and temperature. The JH mimic had, however, no significant effect on the duration of megalopa development. A significant synergism between methoprene, salinity and temperature was not observed. It can be concluded from the results that methoprene does not inhibit metamophosis of R. harrisii larvae at the 0.1 ppm level or lower. Reduction in survival of zoeal stages and increased duration of zoeal development with increasing concentrations of methoprene are presumably related to stress.     

Ultrastructural study of the exoskeleton of the estuarine crab Rhithropanopeus harrisii: Effect of the insect growth regulator Dimilin® (diflubenzuron) on the formation of the larval cuticle

Ultrastructure of larval cuticle during the molt cycle of the estuarine crab Rhitropanopeus harrisii (Gould) (Crustacea: Brachyura) was studied in control larvae as well as in larvae exposed to 10 ppb of the insect growth regulator Dimilin^® (diflubenzuron). First zoeal larvae were used as test organisms. It has earlier been shown that 10 ppb Dimilin is lethal to zoeal larvae of R. harrisii, and nearly all exposed larvae died during molting to the next stage (Christiansen et al., 1978). Distinct differences in structure of the cuticle were found between the two groups of larvae. Both endocuticle and exocuticle appear to be deformed in Dimilin-treated larvae, whereas formation of epicuticle did not seem to be affected. The results indicate that Dimilin probably inhibits chitin synthesis in crab larvae as shown earlier by several authors for insect larvae.     

Distribution and structure of benthic assemblages in Puget Sound,Washington, USA

Benthic infauna was collected with a 0.2 m² van Veen grab at 48 stations in Puget Sound, Washington, USA during February–March 1969. All the crustaceans, lamellibranchs, and echinoderms were identified and counted. Particle size distributions and nitrogen contents of the sediments, depths, and temperature and salinity of the bottom water were determined at all stations. The first three factors of a factor analysis applied to between-stations measures of affinity, explained 41.26% of the total variance. The first factor had representative stations on shallow-water mud bottoms, the second factor on bottoms dominated by coarse sediments, and the third factor had representative stations on deep-water mud bottoms. The multiple correlation coefficients for each of the three factors with depth and mean particle size of the sediments were 0.752, 0.897, and 0.706, respectively. The factor analysis did not result in clusters of stations that could be interpreted as discrete benthic communities. The number of species per 0.6 m² ranged from 8 to 55 species, with a mean of 26.4 species. The frequency distribution of specimens among the species was in good agreement with a log-normal distribution. The number of specimens per 0.6 m² ranged from 57 to 1733 specimens, with a mean of 584.0 specimens. The species diversity ranged from 0.34 bits/individual to 4.35 bits/individual, and there was a weak trend of increasing diversity towards coarser sediments. The first three factors applied to the matrix of between-species correlation coefficients explained 43.72% of the total variance. With each factor there was one group of species with high positive loadings and one group with high negative loadings. Each group of species could be identified with particular environments. The standing crop, measured as ash-free dry weight, ranged from 0.490 to 54.093 g/m², with a mean of 13.752 g/m². Only about 24% of the variability in standing crop could be attributed to variability in sediment types, nitrogen content in the sediments, salinity of the bottom water, and depth.