Rates of fission, somatic growth and gonadal development of a fissiparous sea star, Allostichaster insignis, in New Zealand

Seasonal changes in population structure and incidence of fission were measured in intertidal and subtidal populations of Allostichaster insignis, a fissiparous sea star. Population size structure was stable over the course of the 1-year study. Sea stars in the subtidal zone attained greater maximum size (mean arm length, R = 35 mm) than those in the intertidal population (20 mm). Fission rates were greatest among small individuals (R < 20 mm). The frequency of fission ranged from 5 to 32% with peaks in early austral summer in the intertidal zone, and in autumn and winter in the subtidal zone. Sexual reproduction occurred in early spring in sea stars larger than 12 mm. The populations were heavily biased toward males. In the laboratory, A. insignis of three size classes (small, R = 9–13 mm; medium, 19–21 mm; and large, 29–31 mm) were fed mussels ad libitum or starved (not fed macroscopic food) for ∼1 year in a 3 × 2 factorial experiment. Small and medium-sized sea stars divided throughout the experiment and the ramets of most individuals regenerated sufficiently to divide again after 6–9 months. Unfed sea stars did not undergo fission (with one exception), had a higher mortality rate, and did not grow. Small, fed sea stars grew significantly faster than medium-sized or large individuals. At the end of the experiment, the pyloric caeca index (a measure of nutritional condition) was greater in fed than in unfed animals. Gonads (only testes were observed) developed in medium-sized and large, fed sea stars. Our field and laboratory results indicate that asexual reproduction in A. insignis predominantly occurs in small, well-nourished individuals. Ramets grow gradually through repeated fission and regeneration to a size (mean length of regenerating arms, R _r ∼ 20 mm) at which they begin to switch to sexual reproduction as the dominant reproductive mode.     

Population structure, growth and recruitment of the euryalinid brittle-star Astrobrachion constrictum (Echinodermata: Ophiuroidea) in Doubtful Sound, Fiordland, New Zealand

The euryalinid brittle-star (snake star) Astrobrachion constrictum (Farquhar) lives coiled around the branches of black coral (Antipathes fiordensis) colonies. Twenty-two vertical transects, 10 m wide by 30 m deep, were swum in Doubtful Sound over a 2.5 yr period from 1993 to 1995. Numbers, disc diameters and colour morphotypes of brittle-stars inhabiting coral colonies were recorded. 36.3% of the coral colonies >200 mm tall (n = 292) hosted ≥1 Astrobrachion constrictum (range 0 to 12). Overall, the population was patchily distributed on the available coral habitat. The dark red colour morphotype of A. constrictum was most common (87%, n = 279) followed by the yellow, striped, and then spotted varieties. The population was comprised mainly of large (≥10 mm disc diam) individuals, and juveniles were rarely encountered, indicating low rates of recruitment or a high mortality of recruits. Disc-diameter data gathered from this and previous studies indicated that growth in A. constrictum is initially rapid, with individuals reaching a disc diameter of 15 mm in ≃2.5 yr; growth decreases with age, as in other deep-sea ophiuroids. Growth rate within years, however, was not constant, with faster growth in the spring/summer. Maximum size for A. constrictum is reached in ≃8 yr at ˜23 mm disc diam. Anecdotal evidence indicates that A. constrictum may not be confined solely to black coral colonies. Received: 25 September 1996 / Accepted: 16 October 1996     

Population dynamics,reproduction and growth of the Indo-Pacific horned sea star, <Emphasis Type="Italic">Protoreaster nodosus</Emphasis> (Echinodermata; Asteroidea)

The horned sea star (Protoreaster nodosus) is relatively common in the Indo-Pacific region, but there is little information about its biology. This study of the population biology of P. nodosus was carried out in Davao Gulf, The Philippines (7°5′N, 125°45′E) between September 2006 and May 2008. Protoreaster nodosus was found in sand and seagrass dominated habitats at a mean density of 29 specimens per 100 m² and a mean biomass of 7.4 kg per 100 m², whereas a significantly lower density and biomass was found in coral and rock dominated habitats. Adult specimens (mean radius R = 10.0 cm) were found at depths of 0–37 m, whereas juveniles (R < 8 cm) were only found in shallow sandy habitats with abundant seagrass (water depth ≤2 m). Increased gonad weights were found from March to May (spawning period), which coincided with an increasing water temperature and a decreasing salinity. Density and biomass did not change significantly during reproduction, but sea stars avoided intertidal habitats. All specimens with R > 8 cm had well developed gonads and their sex ratio was 1:1. Protoreaster nodosus grew relatively slowly in an enclosure as described by the exponential function G = 7.433 e^{−0.257 × R} . Maturing specimens (R = 6–8 cm) were estimated to have an age of 2–3 years. Specimens with a radius of 10 cm (population mean) were calculated to have an age of 5–6 years, while the maximum age (R = 14 cm) was estimated as 17 years. Potential effects of ornamental collection on the sea star populations are discussed. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Validation of daily increment deposition in otoliths. Age and growth determination of Aphia minuta (Pisces: Gobiidae) from the northwest Mediterranean

The transparent goby Aphia minuta (Risso, 1810) is one of the main target species of the small-scale fishery off the Island of Majorca. Otolith microstructure and length-frequency analysis were used to study the age and growth of this species during the 1982/1983 and 1992/1993 fishing seasons. Daily periodicity of increment formation was determined by experiments with marked otoliths in individuals maintained in captivity. The length range of the catches during the 11 yr period was between 12 and 49 mm, with a main distribution (89%) between 24 and 40 mm. Otolith age-readings indicate that the population exploited in the commercial fishery consists of seven age-groups (2 to 8 mo old), with a very high proportion of individuals (95%) between 3 and 6 mo old. Population growth-curves revealed no differences between males and females. The growth parameters for the whole population are: asymptotic length, L _∞ = 53.69 mm; growth coefficient, K = 2.23 yr⁻¹; theoretical age at length zero, t ₀ = −0.005 yr. Those individuals of A. minuta caught in Majorca during the winter period reached a maximum age of 7 or 8 mo. Received: 30 December 1996 / Accepted: 16 April 1997     

A laboratory study of behavioral interactions of the Antarctic keystone sea star Odontaster validus with three sympatric predatory sea stars

The circumpolar sea star Odontaster validus is ubiquitous in the nearshore marine benthos of Antarctica. Despite its ecological importance, little is known of its behavioral interactions with other common sympatric sea stars. To examine these interactions we employed time-lapse video analyses conducted in a large laboratory tank (1.8 m diameter circular tank, 1,629 L). In each experimental trial, 34 adult O. validus were placed in a tight circular grouping on one side of the tank, and one adult individual of one of three common sympatric species of predatory sea star (Labidiaster annulatus, Diplasterias brandti, or Perknaster aurorae) was placed on the opposite side of the tank. Digital images of sea star movements were then captured at one min intervals over a 24 h period and aspects of sea star movements subsequently analyzed. Each 24 h treatment was replicated three times, as was a control treatment consisting only of O. validus. O. validus had significantly elevated levels of activity in the presence of P. aurorae when compared with the other two sea stars (potential chemically mediated response), and displayed a distinct “flight response” (change in direction and twofold to sixfold increase of speed) upon tactile contact with this species. Moreover, an “alarm response” was detected when individuals of O. validus that encountered a fleeing conspecific also fled the vicinity. In contrast, our results indicated that O. validus displays virtually no chemical or tactile behavioral responses to the large multi-armed L. annulatus and only weak tactile responses to D. brandti.     

Ontogenetic habitat shift,population growth,and burrowing behavior of the Indo-Pacific beach star, <Emphasis Type="Italic">Archaster typicus</Emphasis> (Echinodermata; Asteroidea)

Archaster typicus, a common sea star in Indo-Pacific regions, has been a target for the ornamental trade, even though little is known about its population biology. Spatial and temporal patterns of abundance and size structure of A. typicus were studied in the Davao Gulf, the Philippines (125°42.7′E, 7°0.6′N), from February 2008 to December 2009. Specimens of A. typicus were associated with intertidal mangrove prop roots, seagrass meadows, sandy beaches, and shoals. Among prop roots, specimens were significantly smaller and had highest densities (131 ind. m⁻²) between November and March. High organic matter in sediment and a relatively low predation rate seemed to support juvenile life among mangroves. Size and density analyses provided evidence that individuals gradually move to seagrass, sandy habitats, and shoals as they age. Specimens were significantly larger at a shoal (maximum radius R = 81 mm). New recruits were found between August and November in both 2008 and 2009. Timing of recruitment and population size frequencies confirmed a seasonal reproductive cycle. Juveniles had relatively high growth rates (2–7 mm month⁻¹) and may reach an R of 20–25 mm after 1 year. Growth rates of larger specimens (R > 30 mm) were generally <2 mm month⁻¹. The activity pattern of A. typicus was related to the tidal phase and not to time of day: Specimens moved over the sediment surface during low tides and were burrowed during high tides possibly avoiding predation. This is one of the first studies to document an ontogenetic habitat shift for sea stars and provides new biological information as a basis for management of harvested A. typicus populations.     

Reproduction and distribution of the invasive European fanworm Sabella spallanzanii (Polychaeta: Sabellidae) in Port Phillip Bay, Victoria, Australia

 The European fanworm Sabella spallanzanii (Gmelin, 1791) was recently introduced to Port Phillip Bay and is now a conspicuous component of most benthic communities. Reproduction of the worm was investigated in a population at Queenscliff over a 2 yr period (October 1995 to October 1997) using gonadal histology. The worms are dioecious (sex ratio 1:1, n=250), and attained sexual maturity at ∼50 mm body length. Reproductive periodicity followed a distinct annual cycle, and spawning proceeded through an extended autumn/winter period. Spawning was broadly synchronous between sexes, and coincided with falling seawater temperatures and shorter day-lengths. The females were highly fecund, and >50 000 eggs were probably shed from large females (>300 mm body length) during the annual spawning period. Breeding cycles of S. spallanzanii in Port Phillip Bay are ∼6 mo out of phase with endemic populations located at similar latitudes in the northern hemisphere. The spread of S. spallanzanii within Port Phillip Bay has been monitored by divers on an annual basis since 1994. The most recent dive survey (1998) indicates that S. spallanzanii has extended its range through out the entire 2000 km² embayment, and has invaded most subtidal habitats. Quantitative estimates of S. spallanzanii abundances were highest on pier pylons (12.5 individuals m⁻², 0.5 to 7 m depths). On sediments, estimates were highest at shallow sites (0.3 m⁻², 7 m depth), but numbers declined significantly with depth (0.1 m⁻², 17 to 22 m depth). Mean worm lengths and biomass were, by contrast, significantly higher at intermediate depths (12 to 17 m) than in shallower (7 m) or deeper (22 m) locations. S. spallanzanii demonstrates a clear preference for growth in sheltered, nutrient-enriched waters, so it may not spread from Port Phillip Bay into the adjacent oceanic waters of Bass Strait; however, in view of S. spallanzanii's current high abundance, fecundity and extended spawning periodicity, there is a high risk of future range expansions, mediated by shipping, into other temperate-water ports. Received: 17 November 1998 / Accepted: 6 January 2000     

Feeding aggregations of sea stars (<Emphasis Type="Italic">Asterias</Emphasis> spp. and <Emphasis Type="Italic">Henricia sanguinolenta</Emphasis>) associated with sea urchin (<Emphasis Type="Italic">Strongylocentrotus droebachiensis</Emphasis>) grazing fronts in Nova Scotia

Migrating feeding aggregations (or fronts) of sea urchins can dramatically alter subtidal seascapes by destructively grazing macrophytes. While direct effects of urchin fronts on macrophytes (particularly kelps) are well documented, indirect effects on associated fauna are largely unknown. Secondary aggregations of predators and scavengers form around fronts of Strongylocentrotus droebachiensis in Nova Scotia. We recorded mean densities of the sea stars Asterias spp. (mainly A. rubens) and Henricia sanguinolenta of up to 11.6 and 1.7 individuals 0.25 m⁻² along an urchin front over 1 year. For Asterias, mean density at the front was 7 and 15 times greater than in the kelp bed and adjacent barrens, respectively. There was strong concordance between locations of peak density of urchins and sea stars (Asterias r = 0.98; H. sanguinolenta r = 0.97) along transects across the kelp–barrens interface, indicating that sea star aggregations migrated along with the urchin front at rates of up to 2.5 m per month. Size–frequency distributions suggest that Asterias at the front were drawn from both the barrens (smaller individuals) and the kelp bed (larger individuals). These sea stars fed intensively on mussels on kelp holdfasts and in adjacent patches. Urchin grazing may precipitate aggregations of sea stars and other predators or scavengers by incidentally consuming or damaging mussels and other small invertebrates, and thereby releasing a strong odor cue. Consumption of protective holdfasts and turf algae by urchins could facilitate feeding by these consumers, which may obtain a substantial energy subsidy during destructive grazing events.     

Age and growth of Electrona antarctica (Pisces: Myctophidae), the dominant mesopelagic fish of the Southern Ocean

Numerically and in biomass, the lanternfish Electrona antarctica is the dominant fish in the vast pelagic region of the Southern Ocean bounded on the north by the Antarctic Convergence and in the south by the Antarctic continental shelf. It is an important krill predator, and in turn is important in the diets of flighted and swimming seabirds. Further, it is the southernmost and coldest-dwelling representative of the globally distributed fish family Myctophidae. The present study was undertaken to estimate the species' growth rate and average life span, to incorporate the information in a basic energy budget, and to compare the growth of E. antarctica with more northerly confamilials. Fishes were aged using primary growth increments that were resolved on sagittal otoliths using three sequential techniques: thin-section grinding and polishing, etching, and scanning electron microscopy (SEM). Based on increment width (0.8 to 1.2 μm), continuity, and previous studies on confamilials, the microincrements were assumed to be deposited on a daily basis. Montages of SEM photomicrographs were constructed for each sagitta to allow the daily rings to be counted over the entire life span of 31 individuals representing the entire size range of the species. Results suggest a larval stage of 30 to 47 d and a maximum life span of 3.5 yr, with females growing faster than males in the last 1.5 yr of life and reaching a larger maximum size. Construction of a simple energy budget using the best information available suggests that a surplus of energy is available to support the observed growth rates (0.05 to 0.07 mm d⁻¹). The results of the present study contrast markedly with previous estimates of an 8 to 11 yr maximum age for E. antarctica. These results provide important data addressing the ecology and population dynamics of the pelagic Antarctic ecosystem. E. antarctica is the end-member species in the continuum of vertically migrating myctophids that extend from the equator to the polar circle. Its growth rate is consonant with that of all other myctophid species examined using primary growth increments to determine age. The present study, in conjunction with earlier studies, suggests that growth rates of mesopelagic species are far higher than previously thought. Received: 12 September 1997 / Accepted: 25 July 1998     

Sea anemones and brittle stars: unexpected predatory interactions during induced in situ oxygen crises

During oxygen crises, benthic faunas exhibit a series of behavioural patterns that reflect the duration and severity of the event. During artificially induced oxygen deficiencies at 24 m depth in the Northern Adriatic Sea, we photographically documented predation by the sea anemones Cereus pedunculatus (Pennant, 1777) and Calliactis parasitica (Couch, 1842) on the brittle star Ophiothrix quinquemaculata (DelleChiaje, 1828). Five predatory events were recorded with four anemones during nine deployments totalling 817 h of observation. Under near-anoxic conditions, individuals of both actinians made contact with, pulled in and consumed the brittle stars. The duration of each predatory event was 1.5–7.5 h. In three of the five events, brittle star remains were regurgitated after an additional 2.0–12.5 h of digestion by the anemones. Our time-lapse sequences demonstrate that oxygen deficiency, beyond eliciting a series of specific behaviours in members of each species, also promotes previously unobserved interspecific interactions. Our results show that sea anemones are not only highly resistant to anoxia, but may also benefit by taking advantage of prey that are more vulnerable to anoxic conditions.     

Growth patterns in oysters, Crassostrea virginica, from different estuaries

We analyzed a data set collected over 15 yr, containing growth data from strains of eastern oysters, Crassostrea virginica (Gmelin, 1791), initiated from parent populations in Long Island Sound, Delaware Bay, and lower Chesapeake Bay. The long-term growth data proved to be a powerful tool for examining patterns of growth differentiation among separated populations of C. virginica. The oyster strains had been grown in a common environment in lower Delaware Bay for up to seven generations. We found that the oyster strains with origins in Long Island Sound were significantly larger over several generations than oyster strains from Delaware Bay and Chesapeake Bay. Chesapeake Bay oyster strains were larger than Delaware Bay oyster strains at 1.5 yr old, but Delaware Bay oysters were larger at 2.5 yr. Year-to-year variation in environmental conditions had a strong significant effect on absolute oyster size and the relative sizes of the oyster strains. Persistent differences between oyster strains from different origins over several generations support a hypothesis that these estuarine populations have experienced long-term genetically-based population differentiation. This result is consistent with hypotheses of population differentiation of oysters based on observations of local reproductive timing. Received: 12 August 1997 / Accepted: 26 May 1998     

Fluoride in Antarctic marine crustaceans

The concentration of fluoride in the body parts of a range of Antarctic crustaceans from a variety of habits was examined with the aim of determining whether fluoride concentration is related to lifestyle or phylogenetic grouping. Euphausiids had the highest overall fluoride concentrations of a range of Antarctic marine crustaceans examined; levels of up to 5477 μg g⁻¹ were found in the exoskeleton of Euphausia crystallorophias. Copepods had the lowest fluoride levels (0.87 μg g⁻¹ whole-body); some amphipods and mysids also exhibited relatively high fluoride levels. There was no apparent relationship between the lifestyle of the crustaceans and their fluoride level; benthic and pelagic species exhibited both high and low fluoride levels. Fluoride was concentrated in the exoskeleton, but not evenly distributed through it; the exoskeleton of the head, carapace and abdomen contained the highest concentrations of fluoride, followed by the feeding basket and pleopods, and the eyes. The mouthparts of E.␣superba contained almost 13 000 μg F g⁻¹ dry wt. Antarctic krill tail muscle had low levels of fluoride. After long-term (1 to 5 yr) storage in formalin, fluoride was almost completely lost from whole euphausiids. Received: 1 April 1998 / Accepted: 29 July 1998     

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     

Size and estimated age of genets in eelgrass, Zostera marina, assessed with microsatellite markers

We examined the spatial distribution of genotypes in a perennial population of eelgrass, Zostera marina L., at two spatial scales. We mapped and sampled 80 ramets in a subtidal area of 20 × 80 m, and an additional 15 ramets in two 1-m² sub-quadrats. Ramets were genotyped for seven polymorphic microsatellite loci. Among a total number of 54 genotypes detected, 12 occurred more than once. The ramets of ten of these genotypes occurred in clusters and represented genets based on their expected multi-locus genotype frequencies. The size distribution of genets was uneven with estimated ramet numbers ranging from 2 to 5000. Whereas some areas displayed a high genet diversity, which may indicate past disturbances, large genets (≥10 m²) predominated in other locations of the sampled plot. Spatial heterogeneity in clone distribution was also obvious at the smaller sampling scale (15 ramets sampled within 1 m²) where the clonal diversity (genets identified/ramets sampled) was 0.24 in one quadrat, and 0.077 in the other. Ramets belonging to the largest clone were maximally 17 m apart, which corresponds to a genet age of 67 yr based on annual rhizome growth rates. We conclude that the spatial arrangement of clones in seagrasses allows inferences about the past demography and the disturbance regime at a given site which may prove useful for coastal zone management of ecologically valuable seagrass meadows. Received: 15 September 1998 / Accepted: 14 November 1998     

Growth of the scyllarid lobsters Ibacus peronii and I. chacei

The growth rates of the morphologically similar scyllarid lobsters Ibacus peronii (Leach, 1815) and I. chacei (Brown and Holthuis, 1998) are described using data from a tag/recapture study and from tagged lobsters kept in captivity. Within particular size classes, we found no differences in moult increments between male and female I. peronii nor between male and female I. chacei. Small individuals of both species always had larger moult increments than larger individuals. For I. peronii, females moulted more frequently than males, and smaller size classes moulted more frequently than larger size classes. Female I. peronii therefore grew more quickly than males and reached their estimated size at sexual maturity (51 mm carapace length) after ∼2 yr. Moulting of I. peronii was seasonal, with most lobsters (96.3%) moulting between October and January. We found no differences in growth rates of I. peronii at two locations along the east coast of Australia: Coffs Harbour in New South Wales (30°18′S; 153°08′E), and Lakes Entrance in Victoria (37°53′S; 148°00′E). For I. chacei, we found no differences in the frequency of moulting between males and females and, because we also found no differences in the moult increments between males and females, the growth rates of both sexes were the same. Received: 14 August 1999 / Accepted: 20 January 2000     

Alpha-male paternity in elephant seals

The aim of this study was to assess paternity of males that dominated mating in harems at northern (Mirounga angustirostris) and southern (M. leonina) elephant seal rookeries using DNA fingerprinting and microsatellite DNA analysis. Southern alpha males had greater reproductive success than most northern alphas at similar-sized harems. Comparison of the relatedness between pups within harems also suggested that fewer males achieved matings in the southern elephant seal population. This was consistent with behavioral observations that suggest greater competition for mates in northern elephant seal harems. Reproductive success was consistent with estimates of mating success in some cases, but lower than expected for some northern elephant seal alpha males. A lower reproductive success than predicted from mating behavior may arise from a variety of factors including sperm competition, male sperm depletion from frequent mating, or reduced fertility. The alternatives are discussed in the context of environmental and historical factors. Received: 17 August 1998 / Received in revised form: 28 March 1999 / Accepted: 30 May 1999     

Light and carbon balance in the seagrass Thalassia testudinum : evaluation of current production models

The production dynamics and carbon balance of Thalassia testudinum in the lower Laguna Madre, Texas, USA, were examined during the 1995 summer period based on in situ photosynthesis vs irradiance (PI) measurements and continuous measurements of underwater photon-flux density (PFD). The validity of applying the H _sat model, used to calculate production for Zostera marina as the product of the maximum rate of photosynthesis (P _max) and daily hours of saturating irradiance (H _sat) was assessed for T. testudinum by comparison with integrated production estimates derived through numerical integration. Gross integrated production values were combined with dark-respiration measurements of photosynthetic (PS) and non-photosynthetic (NPS) tissues and areal biomass to generate daily whole-plant carbon balance. Production and whole-plant carbon balance are discussed in relation to surface and underwater PFD measurements, biomass and other physical and chemical parameters collected during a 1 yr period from January to December 1995. The H _sat model significantly underestimated production during all summer months, averaging 70% of integrated production over the entire study period. Gross integrated production ranged between 11.5 mg C g⁻¹ leaf dry wt d⁻¹ in June (during a period of unseasonably low PFDs caused by a drift-alga mat covering the seagrass bed) to 26.7 mg C g⁻¹ leaf dry wt d⁻¹ in July. Modeled net carbon gain was highest in July at 454 mg C m⁻² d⁻¹ (1.4 g dry wt m⁻² d⁻¹), sufficient to account for measured rates of leaf production in the study area and representative of T. testudinum populations of low productivity. During part of the summer period, however, the population was in negative carbon balance. The relatively low productivity of this population and the periods of negative carbon balance are attributed to low net photosynthesis:dark respiration (P _net:R _d) ratios, sporadic low-light periods, the small fraction of PS tissue relative to whole-plant biomass (5 to 13%) and nutrient limitation. Production models are sensitive to both light availability and the proportion of PS tissue supporting NPS biomass as reflected in whole-plant P _net:R _d ratios. Received: 13 August 1997 / Accepted: 6 March 1998     

Seasonal vertical distribution and population dynamics of the chaetognath Parasagitta elegans in the water column and hyperbenthic zone of Conception Bay, Newfoundland

The vertical distribution and population dynamics of the chaetognath Parasagitta elegans Verrill were determined in the water column and hyperbenthic zone of Conception Bay, Newfoundland from April 1997 to June 1998. The water column depth at the study site (47°32.2′N; 53°07.9′W) was 235 m. The temperature below the thermocline was <0 °C the year round. Chaetognath samples from the water column were collected with a Tucker Trawl. Those from the hyperbenthic zone, were collected with an epibenthic sledge. Depending upon whether the hyperbenthic zone was assumed to extend either 1 m or 10 m above bottom, the grand mean, areal abundance of chaetognaths in the hyperbenthic zone ranged from 6% to 40% of the total abundance in the water column (including the hyperbenthic zone), and the grand mean, areal biomass ranged from 25% to 77%. Large, mature individuals were collected only in the hyperbenthic zone, whereas small, immature individuals were collected primarily in the water column. According to body length and ovary maturity data, three cohorts were identified in the hyperbenthic zone during the study period. Within each cohort, the length frequency of reproductively mature individuals was bimodal, with groups of mean length 33 mm and 41 mm reproducing from May to October. The recruitment period of juvenile chaetognaths extended from July to February, coinciding with the recruitment period of copepods. The estimated individual growth rate of P. elegans was 1.0 mg C year⁻¹. The approximate generation time of the two groups of individuals with mean length at maturity of 33 mm and 41 mm was 450 and 780 days, respectively. This study demonstrates that a failure to sample the large, mature P. elegans living in the hyperbenthic zone leads to serious underestimates of the total abundance and biomass of chaetognaths and an inaccurate picture of seasonal population dynamics. Received: 8 September 1999 / Accepted: 15 September 2000     

The amphipod Corophium multisetosum (Corophiidae) in Ria de Aveiro (NW Portugal). II. Abundance, biomass and production

 The abundance and biomass of Corophium multisetosum Stock, 1952 were determined from benthic corer samples collected monthly over 1 yr in the upper reaches of Canal de Mira (Ria de Aveiro, Portugal). Both density and biomass over the sampling period were negatively correlated with water temperature and positively correlated with chlorophyll a concentration in the sediment. C. multisetosum density was significantly negatively correlated with plant biomass and positively correlated with salinity. The nature of the sediment, favourable environmental conditions, high availability of food and low interspecific competition allowed the population to reach a maximal density of 200 × 10³ individuals m⁻² and a maximal biomass (ash-free dry wt, AFDW) of 62 g_AFDW m⁻². The population was highly productive, especially during the autumn/winter period. Production, estimated by two different methods (Hynes method: 251 g_AFDW m⁻² yr⁻¹; Morin–Bourassa method: 308 g_DW m⁻² yr⁻¹), was much higher than the values reported for other Corophium species. The annual P:Bˉ ratio (10) was high, but similar to values reported for Swedish populations of C. volutator and lower than the values estimated from Mediterranean populations of C. insidiosum. Received: 8 October 1999 / Accepted: 22 June 2000     

Habituation in the green frog, Rana clamitans

Habituation and its role in the dear enemy effect was investigated in a population of green frogs, Rana clamitans. Green frogs have a prolonged breeding season, and males defend territories centered around suitable oviposition sites. We tested the prediction that male green frogs will habituate to broadcasts of synthetic conspecific stimuli. Our results indicate that male green frogs can discriminate familiar from unfamiliar stimuli. We suggest that habituation helps to mediate the territorial interactions between male green frogs. Strangers present a greater threat than familiar neighbors. By habituating to the advertisement vocalizations of their near neighbors, males avoid costly interactions with individuals that are not a major threat to their territories. Received: 20 July 1998 / Accepted after revision: 6 September 1998