Population biology of Dyopedos monacanthus (Crustacea: Amphipoda) on estuarine soft-bottoms: importance of extended parental care and pelagic movements

The suspension-feeding amphipod Dyopedos monacanthus (Metzger, 1875) is a common epibenthic amphipod that lives on self-constructed “mud whips” (built from filamentous algae, detritus and sediment particles) in estuaries of the northern North Atlantic Ocean. The population biology of D. monacanthus at a shallow subtidal site in the Damariscotta River Estuary (Maine, USA) was examined between July 1995 and July 1997. The resident population at the study site was dominated by adult females during most months of the year. High percentages of subadults were found in late summer/early fall. Often, between 10 and 20% of the adult females were paired with males, and the percentage of ovigerous females varied between 40 and 100%, indicating continuous reproduction. The percentage of parental females varied between 40 and 80% during most months, but dropped to levels below 20% during summer/early fall. The average size of amphipods on their own mud whips was ∼4 mm during the summer/early fall, after which it increased continuously to >7.0 mm in March or April, and then dropped again. In March and April, the average number of eggs and juveniles female⁻¹ was ∼100 eggs and 55 juveniles, while during the summer/early fall the average number of eggs female⁻¹ was <20 and that of juveniles female⁻¹ was <10. Many juveniles grew to large sizes (>1.4 mm) on their mothers' whips in winter/early spring but not in the summer/fall. The average number of amphipods at the study site was low in late summer/early fall (<50 individuals m⁻²), increased steadily during the winter, and reached peak densities of >3000 individuals m⁻² in April 1996 (>1600 individuals m⁻² in May 1997), after which densities decreased again. The decrease of the D.␣monacanthus population at the study site coincided with a strong increase of amphipods found pelagic in the water column. This behavioural shift occurred when temperatures increased and benthic predators became more abundant and active on shallow soft-bottoms, suggesting that D. monacanthus at the study site is strongly affected by predation. The effects are direct (by predation on amphipods) and indirect (by reducing duration of extended parental care and enhancing pelagic movements). Both extended parental care and pelagic movements are important behavioural traits of D.␣monacanthus (and other marine amphipods), and significantly affect its population dynamics. Received: 18 January 1998 / Accepted: 27 May 1998     

Reproductive biology of a deposit-feeding amphipod, Casco bigelowi, with extended parental care

The reproductive traits of a deposit-feeding amphipod that engages in extended parental care were examined. At the study site in Lowes Cove, Maine, USA, Casco bigelowi (Blake, 1929) occurred in highest densities in soft sediments just below mean low water (MLW). During most months, the sex ratio was ≃1. Many females hosted males in their burrows throughout the summer, but after fertilization of females in September, all adult males disappeared from the study area. In October almost 80% of the females were ovigerous, and in November about the same percentage was parental, i.e. caring for juveniles in their burrows. The females produced only one brood each in late fall which they accommodated in their burrows for 2 mo or longer. The average number of juveniles per female was ∼20 in November, and continuously decreased until January. Juveniles reached sizes >10 mm length in the maternal burrows. In early December the first juveniles were found in their own burrows, but major recruitment took place in late December and January. It is concluded that for C. bigelowi, the delay of recruitment into the winter months with low predation pressure and the large offspring size at this time are major advantages gained by extended parental care. C. bigelowi is host to the peritrich ciliate Cothurnia sp. on its gills, and during the summer months >70% of all amphipods had ciliates on their gills. Juveniles still living in their mother's burrows showed infestation rates similar to that of the parent; those of highly infested mothers were more heavily infested than those of “clean” mothers. Facilitated epibiont transmission during intimate and long-lasting (2␣mo) parent–offspring associations may be a consequence of extended parental care. Received: 25 November 1997 / Accepted: 14 April 1998     

Reproductive biology of a wood-boring isopod, Sphaeroma terebrans, with extended parental care

The isopod Sphaeroma terebrans Bate, 1866 burrows in aerial roots of the red mangrove Rhizophora mangle L. The burrows serve as shelter and as a reproductive habitat, and females are known to host their offspring in their burrows. I examined the reproductive biology of S. terebrans in the Indian River Lagoon, a shallow lagoon stretching for ∼200 km along the Atlantic coast of Florida, USA. Reproductive isopods were found throughout the year, but reproductive activity was highest in the fall and during late spring/early summer. During the latter periods, large numbers of subadults established their own burrows in aerial roots. The average numbers of S. terebrans per root were high during the fall, but decreased during the winter and reached lowest levels at the end of the summer. Females reached maturity at a larger size than males, but also grew to larger sizes than the males. The average size of females varied between 8 and 10 mm, the average size of males between 6.5 and 8.5 mm. The number of embryos female⁻¹ was strongly correlated with female body length. No indication for embryo mortality during development was found. Parental females (i.e. with juveniles in their burrows) hosted on average between 5 and 20 juveniles in their burrows (range 1 to 59). Most juveniles found in female burrows were in the manca stage and 2 to 3 mm in body length. Juveniles did not increase in size while in the maternal burrow, and juveniles of similar sizes could also be found in their own burrows. Males did not participate in extended parental care, since most of them left the females after copulation. Many females that were born in the summer produced one brood in the fall and a second during winter/early spring. Females that were born in the fall produced one brood during spring/early summer, but then probably died. Extended parental care in S. terebrans is short compared to other peracarid crustaceans. It is concluded that this reproductive strategy in S. terebrans serves primarily to shelter small juveniles immediately after they emerge from the female body, when their exoskeleton is still hardening and their physiological capabilities are still developing. Thus, in S. terebrans, extended parental care probably aids to protect small juveniles from adverse physical conditions in their subtropical intertidal habitat. Received: 9 December 1998 / Accepted: 24 June 1999     

Some ecological observations on fishes associated with drifting seaweed off Tohoku coast,Japan

Fishes associated with drifting seaweed were sampled in the Tohoku area, northwest Pacific, in late spring 1988, to analyse the community structure and ecological relationships of the dominant species.Seriola quinqueradiata Temminck et Schlegel was the dominant species and its escort was composed ofThamnaconus (Navodon)modestus, Sebastes inermis, Hyperoglyphe japonica andEnedrias nebulosus. E. nebulosus, which fed essentially on Gammarid crustaceans, was collected mainly in Sendai Bay; the other species, found in offshore water, were plankton feeders and did not exhibit any strong competition, indicating close cohabitation. The abundance of drifting seaweed in the Tohoku area peaked in May–June and decreased in July; they seemed to come from the nearest coast. JuvenileS. quinqueradiata (> 15 cm long) were collected with set nets in Sendai Bay during summer. They seemed to use drifting seaweed (along with the warm Kuroshio current) as a means of transportation during their migration from the southern spawning ground to northern Tohoku area. Drifting seaweed could be considered as a nursery, and the association of fishes with floating algae might have an influence on species recruitment.     

Reproductive biology of a filter-feeding amphipod, Leptocheirus pinguis , with extended parental care

Leptocheirus pinguis (Stimpson, 1853) is a widely distributed, abundant, endobenthic amphipod that engages in extended parental care, i.e. females host their juveniles in their burrows for extended time periods. I examined reproduction and population biology of L. pinguis at mean low water (MLW) in muddy sediment in Lowes Cove, Maine, USA. Cores around individual burrows were taken monthly in 1994 and 1995, and four seasonal samples were taken at different tidal heights. During the major reproductive periods in spring/early summer and in the fall, females produced several consecutive broods and hosted growing offspring in their burrows. Juveniles remained in their mothers' burrows until they reached a length of 5 mm (approximately one-third adult size) or more. At the study site, the majority of amphipods in individual burrows were adult females. Following the main reproductive periods, subadult individuals were found in their own burrows, but densities did not increase following the reproductive period in spring/early summer 1994, probably because large numbers of L. pinguis emigrated via the water column between June and December. L. pinguis is an annual species. Many members of the cohort born in spring/early summer start reproducing in the fall, and survive until the following spring when they produce several broods. Members of the cohort born in the fall start reproducing the following spring and also produce several consecutive broods. Both the spring/early summer and fall cohorts die off after the major reproductive period in the following spring/early summer. High standing stocks of microphytobenthos occur in soft-bottoms at MLW, and I conclude that L. pinguis can engage in extended parental care there because its food is abundant year-round. The limited expandability and low stability of burrows in soft-bottoms at MLW do not permit long persistence of parent–offspring groups in L.␣pinguis. Received: 27 March 1997 / Accepted: 30 July 1997     

Feeding and predation impact of two chaetognath species, Eukrohnia hamata and Sagitta gazellae, in the vicinity of Marion Island (southern ocean)

The predation impact of the two chaetognaths Eukrohnia hamata and Sagitta gazellae on mesozooplankton standing stock were investigated in three depth layers during two 24 h stations occupied in the vicinity of Marion Island in late austral summer (April/May) 1986. The zooplankton community at both stations was dominated by small copepods (Oithona spp., Microcalanus spp.), which accounted for >95% of total zooplankton abundance. Chaetognaths comprised <2% of total zooplankton abundance. E. hamata constituted >95% of the total chaetognath stock. The general trend in both species was decreasing abundance with increasing depth, which appeared to be correlated to the distribution of copepods (r ² = 0.45; P <0.05). Gut-content analysis showed that copepods (mainly Oithona spp., Calanus spp. and Rhincalanus gigas) and ostracods were the main prey of both species, accounting for 87 and 61% of the total number of prey in E. hamata and S.␣gazellae stomachs, respectively. In the guts of S.␣gazellae, pteropods (Limacina spp.) and chaetognaths were also well represented. The mean number of prey items (NPC) for E. hamata ranged from 0.02 to 0.06 prey individual⁻¹ which corresponds to an individual feeding rate (Fr) of between 0.05 and 0.12 prey d⁻¹. For S.␣gazellae, the NPC values were higher, varying between 0.04␣and 0.20 prey individual⁻¹, or between 0.15 and 0.76 prey d⁻¹. The daily predation impact of the two chaetognaths was estimated at between 0.3 and 1.2% of the copepod standing stock or between 7 and 16% of the daily copepod production. Predation by S. gazellae on chaetognaths accounted for up to 1.6% of the chaetognath standing stock per day. Received: 26 November 1996 / Accepted: 31 October 1997     

Effect of development rate on the swimming, escape responses, and morphology of yolk-sac stage larval American plaice, Hippoglossoides platessoides

To examine the impact of development rate on swimming performance, escape response, and morphology, yolk-sac larvae of American plaice (Hippoglossoides platessoides, Fabricius) were reared at two temperatures (5 and 10 °C). Videomicroscopy and silhouette collimation videography were used to examine swimming, escape behaviour, and morphology (standard length, finfold area, and yolk-sac area) of individual larvae. Larvae were examined from 0 d post hatch (dph) to 14 dph for the 5 °C treatment group and from 0 to 6 dph for the 10 °C treatment group (3 August to 17 August 1996). Since larvae were not fed, yolk-sac reserves were essentially exhausted by 14 and 6 dph for the 5 and 10 °C treatment groups, respectively. To control for the effect of testing temperature on behaviour, larvae from each temperature treatment were tested at both 5 and 10 °C. Testing temperature had an effect on some swimming parameters but not on escape response. Swimming performance, escape response, and morphology varied with age, while only morphology and escape response varied with development rate. Morphology and swimming performance, and morphology and escape response were found to be correlated as determined by canonical correlation. This study suggests that both types of swimming behaviours should be examined when developing models of the impacts of predation on the early life history of larval fish. Received: 13 September 1999 / Accepted: 21 June 2000     

Oceanic diffusion and the pelagic insects Halobates spp. (Gerridae: Hemiptera)

Five pelagic Halobates species occupy a vast area from 40^∘ north to 40^∘ south in the three major oceans. Oceanic diffusion, constantly acting to disperse these insects, must be an important factor in determining their life history and distribution. We investigated the effects of oceanic diffusion on the following aspects of these insects. (1) The estimated radius of a patch of Halobates could be expanded by oceanic diffusion alone from an initial point of origin to 1250 km in 60 d. This distance is about 1/12 of the maximum distributional range of H. micans in the Pacific Ocean. Mutual encounter rates due to oceanic turbulence could be as high as 11 d⁻¹ even at low population densities (100 ind km⁻²). This suggests that individuals from their original habitat could find mates even when they had been carried a long distance. Thus, extensive gene mixing may occur over the whole range of a species' distribution. (2)␣Estimated growth rates are rather low (0.0026 to 0.0079 d⁻¹) compared with those of other insects. However, they are offset by a long life span (over 90 d) and an extended oviposition period (perhaps over 2␣months). Thus, pelagic Halobates spp. appear to have adopted a strategy of slow growth and prolonged longevity to cope with living in an unstable physical environment that is constantly disturbed by storms and winds. Received: 5 February 1995 / Accepted: 30 October 1997     

Temporal succession and spatial segregation of clupeoid larvae in the coastal waters off the Tanshui River Estuary, northern Taiwan

In the coastal waters off the Tanshui River Estuary in northern Taiwan, eight species of clupeoid larvae were observed. They exhibited a distinct temporal succession in association with seasonal temperature changing. The time of peak abundance of Etrumeus teres was in January, Engraulis japonicus in late April, Sardinella spp. in mid-May, Dussumieria elopsoides in early June, Thryssa dussumieri in late June, Stolephorus insularis in mid-September, Encrasicholina heteroloba in early October and E. punctifer in mid-November. The time intervals of the temporal succession of the fishes were approximately 15 to 25 d in the spring/summer and 25 to 35 d in the autumn/winter fishing seasons. Also, they showed spatial segregation by distributing in areas with different water depths: Sardinella spp. at a water depth of 10 to 20 m, T. dussumieri at less than 10 m, E. heteroloba at 20 to 50 m and E. punctifer at 10 to 40 m. The larvae of these sympatric clupeoid species segregated their nursing periods and areas apparently to reduce competition for habitat and thus to maximize the utilization of resources. Received: 3 March 1997 / Accepted: 19 March 1997     

In situ swimming and settlement behaviour of larvae of an Indo-Pacific coral-reef fish, the coral trout Plectropomus leopardus (Pisces: Serranidae)

Late larvae of the serranid coral trout Plectropomus leopardus (Lacepède), captured in light traps, were released during the day both in open water and adjacent to two reefs, and their behaviour was observed by divers at Lizard Island, northern Great Barrier Reef. Coral trout larvae (n = 110) were present in light-trap catches from 18 November to 3 December 1997, including new moon (30 November). The swimming speed of larvae in open water or when swimming away from reefs was significantly greater (mean 17.9 cm s⁻¹) than the speed of larvae swimming towards or over reefs (mean 7.2 cm s⁻¹). Near reefs, larvae swam at average depths of 2.7 to 4.2 m, avoiding 0 to 2 m. In open water, swimming depth varied with location: larvae >1 km east of Lizard Island swam steeply downward to >20 m in 2 to 4 min; larvae >1 km west oscillated between 2.6 and 13 m; larvae 100 to 200 m east of Lizard Island oscillated between 0.8 and 15 m. Nearly all larvae swam directionally in open water and near reefs. In open water, the average swimming direction of all larvae was towards the island, and 80% (4 of 5) swam directionally (p < 0.05, Rayleigh's test). Larvae swam directionally over the reef while looking for settlement sites. The frequency of behaviours by larvae differed between two reefs of different exposure and morphology. Depending on site, 26 to 32% of larvae released adjacent to reefs swam to open water: of these, some initially swam towards or over the reef before swimming offshore. In some cases, offshore-swimming seemed to be due to the presence of predators, but usually no obvious cause was observed. Depending on the reef, 49 to 64% of the larvae settled. Non-predatory reef residents aggressively approached 19% of settlers. Between 5 and 17% of the larvae were eaten while approaching the reef or attempting to settle, primarily by lizardfishes but also by wrasses, groupers and snappers. A higher percentage of larvae settled in the second week of our study than in the first. Average time to settlement was short (138 s ± 33 SE), but some larvae took up to 15 min to settle. Average settlement depth was 7.5 to 9.9 m, and differed between locations. No settlement took place on reef flats or at depths <4.2 m. Larvae did not appear to be selective about settlement substrate, but settled most frequently on live and dead hard coral. Late-stage larvae of coral trout are capable swimmers with considerable control over speed, depth and direction. Habitat selection, avoidance of predators and settlement seem to rely on vision. Received: 7 July 1998 / Accepted: 26 January 1999     

Low molecular-weight factor from Plesiastrea versipora (Scleractinia) that modifies release and glycerol metabolism of isolated symbiotic algae

When symbiotic dinoflagellate algae (Symbiodinium sp., isolated from the coral Plesiastrea versipora) were incubated with NaH¹⁴CO₃ in the light in seawater, they released 22.69±9.16 nmol carbon/10⁶ algae. Release of photosynthetically fixed carbon was stimulated more than six-fold for algae incubated in host-tissue homogenate (148.54±97.03 nmol C/10⁶ algae) and more than four-fold (102.00±49.16 nmol C/10⁶ algae) for algae incubated in a low molecular weight fraction (≤1 000 M _r ) prepared from host homogenate. Soluble released ¹⁴C-labelled products, as determined by chromatography and autoradiography, were the same when algae were incubated in either host homogenate or the low molecular weight fraction. After 4 h incubation in the light (300 mol photons m⁻² s⁻¹),␣intracellular␣glycerol increased in algae incubated with the low molecular weight fraction (an increase of 0.39 to␣0.67 nmol glycerol/10⁶ algae) compared with little or no increase in algae incubated in seawater (0 to 0.12 nmol glycerol/10⁶ algae). Partial inhibition of triglyceride synthesis (up to 51%) was also observed when algae were incubated in the low molecular weight fraction. All these effects are the same as those observed when algae were incubated in host homogenate. These data indicate that the “host release-factor” activity of P.␣versipora is a compound of low molecular weight. Received: 13 February 1997 / Accepted: 24 October 1997     

Behaviour and tolerance to hypoxia in juvenile Norway lobster (Nephrops norvegicus ) of different ages

The annual occurrence of hypoxia (<25% oxygen saturation) in the bottom waters along the Swedish west coast coincides with the postlarval settlement of Norway lobster, Nephrops norvegicus (L.). This study investigates behaviour and the experimental effects of low oxygen concentrations in juvenile N. norvegicus of different ages. All experimental individuals were reared to the juvenile (postlarval) stage in the laboratory and then given sediment as a substratum. Behavioural responses to low oxygen concentrations were tested in early and late Postlarvae 1 exposed to normoxia (>80% oxygen saturation, pO₂ > 16.7 kPa), moderate hypoxia (30% oxygen saturation, pO₂ = 6.3 kPa) and hypoxia (25% oxygen saturation, pO₂ = 5.2 kPa). The experiments were run for a maximum period of 24 h or until individuals died. Behaviour was studied using sequential video recordings of four behavioural activities: digging, walking, inactivity or flight (escape swimming up into the water column). Behaviour and mortality changed with lowered oxygen concentrations; energetically costly activities (such as walking) were reduced, and activity in general declined. In normoxia, juveniles initially walked and then burrowed, but when exposed to hypoxia they were mainly inactive with occasional outbursts of escape swimming. To increase oxygen availability the juveniles were observed to raise their bodies on stilted legs (similar to adults in hypoxic conditions), but oxygen saturations of 25% were lethal within 24 h. The results suggest that the main gas exchanges of early postlarval stages occur over the general body surface. Burrowing behaviour was tested in Postlarvae 1 and 2 of different ages held in >80% oxygen saturation for 1 wk. The difference in time taken to complete a V-shaped depression or a U-shaped burrow was measured. The results showed a strong negative relationship between postlarval age and burrowing time, but all individuals made a burrow. Juveniles were more sensitive to hypoxia than adults. Thus, the possible consequences of episodic hypoxia for the recruitment of Nephrops norvegicus and for the recolonization of severely affected areas are discussed. Received: 4 August 1996 / Accepted: 11 October 1996     

Effect of an organophosphorus insecticide, fenitrothion, on survival and osmoregulation of various developmental stages of the shrimp Penaeus japonicus (Crustacea: Decapoda)

The toxicity of fenitrothion was determined in larvae (nauplii, Zoeae 1 to 3, Mysis 1 to 3), postlarvae (PL stages) and juvenile shrimp (Penaeus japonicus Bate), in two media, seawater (SW) and diluted seawater (DSW) (1100 and 550 mosM kg⁻¹, ≃ 37 and 19‰ S). The effects of fenitrothion on the osmoregulatory capacities (OC) of juveniles were recorded. A gill and epipodite histopathological study was also conducted. For larvae in seawater, 24 and 48 h LC₅₀s ranged from 32.9 μg l⁻¹ (Zoeae 2) to 10.7 μg l⁻¹ (Mysis 3), and from 3.9 μg l⁻¹ (Zoeae 3) to 2.0 μg l⁻¹ (Mysis 3), respectively; 48 and 96 h  LC₅₀s in postlarvae (PL) at the same salinity ranged from 1.8 μg l⁻¹ (PL1) to 0.6 μg l⁻¹ (PL5), and from 0.3 μg l⁻¹ (PL7) to 0.4 μg l⁻¹ (PL15). In juveniles, 96 h LC₅₀s were 0.8 μg l⁻¹ in seawater and 1.5 μg l⁻¹ in diluted seawater. From hatching to juvenile stages, the overall trend was a rapid decrease (from nauplii to PL5–PL7) followed by a slight increase (from PL7 to PL15 and juveniles) in the shrimp's ability to tolerate the insecticide. In juveniles kept in seawater and in diluted seawater, fenitrothion decreased the osmoregulatory capacity (OC = difference between the hemolymph osmotic pressure and the osmotic pressure of the medium) at both lethal and sublethal concentrations. This effect was time- and dose-dependent. In SW, the decrease in hypo-OC was ˜ 25% at sublethal concentrations and ˜ 35% at the 96 h LC₅₀. In DSW, the decrease in hyper-OC was ˜ 10 to 15% at sublethal concentrations. In SW, shrimp were able to recover their OC in less than 48 h when transferred to water free of pesticide. In DSW, recovery at 48 h was only possible after exposure to the lowest tested sublethal concentration. Haemocytic congestions (thrombosis) of the gills, lamellae necrosis and other alterations of gills and epipodites (breakage of the cuticle, reduction of the hemolymph lacunae) were noted in juveniles exposed to lethal and sublethal concentrations of fenitrothion. Received: 7 October 1996 / Accepted: 13 November 1996     

Grazing of phytoplankton by copepods in eastern Antarctic coastal waters

Chlorophyll a, primary productivity and grazing by copepods on phytoplankton were measured in the upper water column during the summer of 1994/1995 at a coastal site near Davis Station, East Antarctica. Chlorophyll a was at a maximum in mid-December, then dropped markedly as the coastal fast ice melted and broke‐out. Phytoplankton biomass increased again from mid‐ to late‐February. Copepods accounted for at least 65% of zooplankton biomass in the water column before sea ice break‐out, whereas larval polychaetes and ctenophores dominated after ice break‐out. Oncaeacurvata was the numerically dominant species throughout the study. The highest grazing rate (8.7 mg C␣m⁻³␣d⁻¹) was recorded on 21 December when O.␣curvata accounted for 64% of the total. Grazing had decreased markedly by 28 December (0.9 mg C m⁻³ d⁻¹); again O. curvata accounted for over 50% of the total ingested. Copepod grazing increased after ice break-out until the last experiment on 20 February (⋍5 mg C␣m⁻³␣d⁻¹). The main species responsible for grazing during this period were O. curvata, Oithonasimilis, Calanoidesacutus and unidentified copepod nauplii. It was estimated that copepods removed between 1 and 5% of primary productivity. Received: 11 October 1996 / Accepted: 22 October 1996     

Age and growth of the mesopelagic squid Ancistrocheirus lesueurii (Oegopsida: Ancistrocheiridae) from the central-east Atlantic based on statolith microstructure

Statolith microstructure was studied in 56 Ancistrocheirus lesueurii (25 to 423 mm of mantle length, ML) caught in the central-east Atlantic. Statolith growth increments were grouped into three main growth zones, distinguished mainly by increment width. The second transition in the statolith microstructure (from Zone 2 to Zone 3) coincides with the life history shift from epipelagic and upper mesopelagic to a bathyal habitat. Second-order bands (mean 27.65 growth increments) and sub-bands (mean 13.6 growth increments) within statolith microstructure appeared to be related to the lunar cycle. Striking sexual dimorphism is reflected in the age and growth rates: males live ca. 1 yr, while females only start maturing at this age and obviously live >1.5 yr. A. lesueurii is a slow growing squid, attaining 25 to 30 mm ML at the age of 100 d. After ontogenetic migrations into bathypelagic waters at ML > 30 to 35 mm, growth rates gradually decrease to the minimum known values for squids. Based on back-calculated hatching dates, A. lesueurii hatches throughout the year with a peak between November and March. Received: 28 August 1996 / Accepted: 31 January 1997     

Role of protein in larval swimming and metamorphosis of Bugula neritina (Bryozoa: Cheilostomatida)

Larvae of the marine cheilostomatid bryozoan Bugula neritina (L.) were prevented from settling for 1, 4 and 8 h by mechanical agitation, following which settlement and metamorphosis success were examined. Settlement rates were significantly affected by swimming time, which decreased from 100% after 2 h to 93.7 ± 4.3% after 8 h. Similarly, metamorphosis to the feeding ancestrula was significantly impaired following a swimming time of 8 h, declining from 93.7 ± 4.3% after 1 h to 65.9 ± 7.0% after 8 h. The resultant colonies grew well for the first 3 wk, following which time, growth patterns became erratic. Growth rate was in all cases highly variable, and did not correlate with enforced swimming times. Larval protein composition was examined after 1, 4 and 8 h swimming time, and post-larval composition 1, 2, 5, 24 and 48 h after settlement using sodium-dodecyl-sulphate polyacrylamide-gel electrophoresis (SDS-PAGE). Individual protein content was measured using a densitometer. Larvae did not consume protein during swimming, however a protein measuring 170 kdaltons was consumed during metamorphosis. These results are discussed in the context of larval settlement and energetics. Received: 19 July 1998 / Accepted: 3 December 1998     

Spawning and larval development of the black turban snail Tegula funebralis (Prosobranchia: Trochidae)

An understanding of spawning and larval development can be fundamental to interpreting the abundance, distribution, and population structure of marine invertebrate taxa. Tegula funebralis (A. Adams, 1855), the black turban snail, has been the focus of numerous ecological studies on the Pacific coast of North America. To date, there are only conflicting and anecdotal reports of spawning, and there is no information on larval or juvenile development for this conspicuous and abundant species. On 19 September 1995, two individuals of T. funebralis were observed free-spawning gametes into seawater in tanks at the Oregon Institute of Marine Biology. Embryos and larvae were subsequently reared to metamorphosis and beyond. Development was pelagic and similar to development described for other trochids, and larvae were observed not to feed at any stage. Larvae began to metamorphose at 5.7 to 6.7 d and settled at 260 μm shell length. Juveniles grew ≃ 10 μm in shell length per day and appeared to feed on detritus. Juveniles lacked some adult diagnostic shell characters, including two columellar nodes and a closed umbilicus. In the field, small (<3 mm) juveniles occurred in the adult habitat on all sampling dates between October and March. Small juveniles were found only under rocks and were most abundant under rocks partially buried in coarse sand, suggesting that juveniles may utilize a specific microhabitat within the adult T. funebralis habitat. Received: 7 October 1996 / Accepted: 17 October 1996     

Feeding, metabolism and growth in the Antarctic limpet, Nacella concinna (Strebel 1908)

Post-prandial increases in metabolism, the specific dynamic action of feeding (SDA), were evaluated in the Antarctic limpet Nacella concinna. O₂ consumption rose to a peak value 2.3 times higher than pre-feeding standard metabolic rates. This peak rise is low for marine ectotherms, but is typical of polar species. There were three peaks in the SDA, the first lasted only for the 1st day, was caused by handling, and was minor. The second was the major peak. It lasted from post-prandial days 4–9 inclusive, and accounted for around 70% of the SDA response. The third peak lasted from day 11 to day 15 and accounted for 30% of the total SDA. A 15-day SDA is much longer than values for temperate species, but is again typical for polar marine ectotherms. NH₃ excretion declined post-prandially from around 0.4 μmol animal⁻¹ h⁻¹ to values between 0.025 and 0.223 μmol animal⁻¹ h⁻¹ throughout the SDA. The total O₂ consumed in the SDA was 90.2 μmol O₂, which converts to 44.7 J of energy. This was 45–50% of the energy consumed in the meal (93.5 J). Pre-feeding O:N ratios, after 26 days without food, were around 1, indicating protein as the sole metabolic substrate prior to initiating the SDA. After feeding, O:N ratios rose to between 2.5 and 19, indicating significant use of lipid or carbohydrate from the food. Experiments were conducted in ambient seawater with enhanced levels of Sr (SrCl added at 800 mg kg⁻¹), and limpets were fed microalgal films also grown in enhanced Sr media. Sr incorporated in the shells during the experiment allowed the measurement of shell increments deposited during the SDA. Between five and eight microgrowth bands were present in the Sr-enhanced increments, which was similar to the number of days in the second SDA peak. The mean shell increment laid down was 17.6 μm. Estimating tissue deposition from measured growth increments and published ash-free dry mass (AFDM) to length relationships produced a value of 0.81 mg AFDM, which converted to 26.4 J of energy, or 25–30% of the energy ingested in the meal. Estimates of growth increments associated with a single SDA have not previously been possible. Overall energy used in the SDA and tissue deposition accounted for 75–80% of the energy ingested; the remainder was probably accounted for by unmeasured costs such as mucus production. Received: 6 June 2000 / Accepted: 20 September 2000     

Life history of Gonatus onyx (Cephalopoda: Teuthoidea): ontogenetic changes in habitat, behavior and physiology

In the present study 670 individuals of Gonatus onyx (Cephalopoda: Teuthoidea) were observed in Monterey Bay, California from a remotely operated vehicle. The vertical distribution of this species was bimodal, with peaks at 400 and 800 m depth during the day and 300 and 500 m during the night. The bimodal distribution reflects a life stage shift between younger, schooling juveniles living in shallower water and older, solitary adults which live deeper. Ontogenetic changes in behavior associated with this life stage shift are reflected in the physiology of the organisms as well. Both aerobic and anaerobic metabolism, as estimated using mitochondrial and glycolytic enzymes, decline with increasing body mass, suggesting reduced locomotory capacity in deeper-living adults. Oxygen consumption rates were also determined in relation to oxygen partial pressure. Oxygen consumption regulation was similar between juvenile and adult squids. The critical oxygen partial pressures (29 to 30 mmHg) correspond precisely to the oxygen concentrations found at the depth of maximal abundance for day and night populations of juveniles and adults, respectively. Behavioral and physiological changes with ontogeny of G. onyx are believed to result from reduced visual predator/prey interactions in the light-limited deep sea. Received: 7 February 1999 / Accepted: 26 January 2000     

Zooplankton responses to hypoxia: behavioral patterns and survival of three species of calanoid copepods

Seasonally recurrent and persistent hypoxic events in semi-enclosed coastal waters are characterized by bottom-water dissolved oxygen (d.o.) concentrations of < 2.0 ml l⁻¹. Shifts in the distribution patterns of zooplankters in association with these events have been documented, but the mechanisms responsible for these shifts have not been investigated. This study assessed interspecific differences in responses to hypoxia by several species of calanoid copepods common off Turkey Point, Florida, USA: Labidocera aestiva (Wheeler) (a summer/fall species), Acartia tonsa (Dana) (a ubiquitous year-round species), and Centropages hamatus (Lilljeborg) (a winter/spring species). Under conditions of moderate to severe hypoxia 24-h survival experiments were conducted for adults and nauplii of these species from August 1994 to October 1995. Experiments on adults used a flow-through system to maintain constant d.o. concentrations. Adults of A. tonsa showed no decline in survival with d.o. as low as 1.0 ml l⁻¹, sharp declines in survival at d.o. = 0.9 to 0.6 ml l⁻¹, and 100% mortality with d.o. = 0.5 ml l⁻¹. Adults of L. aestiva and C. hamatus were more sensitive to oxygen depletion: both species experienced significant decreases in survival for d.o. = 1.0 ml l⁻¹. Nauplii of L. aestiva and A. tonsa showed no significant mortality with d.o. = 1.1 to 1.5 ml␣l⁻¹ and d.o. = 0.24 to 0.5 ml l⁻¹, respectively. In addition, experiments investigating behavioral avoidance of moderate to severe hypoxia were carried out for adults of all three species. None of the three species effectively avoided either severely hypoxic (d.o. < 0.5 ml l⁻¹) or moderately hypoxic (d.o. ≈ 1.0 ml l⁻¹) bottom layers in stratified columns. These results suggest that in␣nearshore areas where development of zones of d.o. < 1.0 ml l⁻¹ may be sudden, widespread, or unpredictable, patterns of reduced copepod abundance in bottom waters may be due primarily to mortality rather than avoidance. Received: 31 August 1996 / Accepted: 24 September 1996