Influence of seasonal variation in temperature, salinity and food availability on module size and colony growth of the estuarine bryozoan Conopeum seurati

Zooid size and colony growth of the estuarine bryozoan Conopeum seurati (Canu) (order: Cheilostomatida; suborder: Malacostegina) were examined over 15 mo at Avonmouth Dock, Avon, England. Data were analysed in conjunction with synchronous measurements of temperature, salinity and food availability. Zooid length, width and area were strongly temperature-dependent, while both food availability and colony growth rate had no significant effect on zooid length, width or area. Salinity and the interaction of temperature and salinity significantly influenced zooid length and area, suggesting that changes in zooid size may result from oxygen limitation in warm waters. The validity of a number of other mechanisms proposed to account for temperature-related changes in zooid size is discussed. The results support the use of zooid size as an indicator of both long-term trends and seasonal variations in temperature in Recent and fossil assemblages as long as data sets are large and the effects of other factors on zooid size are considered. Colony growth rate was found to be significantly influenced by both the amount of food available to the colonies and the combined effect of temperature and food availability, suggesting that growth rate increases as food increases, but that the former may be limited at low temperatures when metabolic rates are low. Received: 30 March 1999 / Accepted: 24 September 1999     

Zooid size parallels contemporaneous oxygen isotopes in a large colony of<Emphasis Type="Italic"> Pentapora foliacea</Emphasis> (Bryozoa)

The relationship between seasonal changes in temperature and the size of zooids within cheilostome bryozoans is explored by comparing zooid size and contemporaneous carbonate mineralogy within a single colony of a perennial species that grew in a highly seasonal environment. Previously published oxygen isotope profile data from two fronds of a large colony of Pentapora foliacea from the Irish Sea record the cyclical patterns related to seasonal changes in temperature experienced by the colony over 3 years of growth. Zooid size data gathered from the same points at which the oxygen isotope data were taken reveal that zooid size and contemporaneous ¹⁸O values covary. These and previously published data show that zooid size profiling can be confidently used to easily estimate rates of growth and longevity of bryozoan colonies, and further illustrate the negative relationship between zooid size and ambient temperature. Some inconsistencies in timing between zooid size and ¹⁸O data may provide insights into the mechanisms behind temperature mediated body size changes in poikilothermic animals.Communicated by P. W. Sammarco, Chauvin     

Seasonality of polypide recycling and sexual reproduction in some erect Antarctic bryozoans

The seasonality of polypide cycling has been investigated for three species of erect bryozoans from Antarctica: Isoseculiflustra rubefacta (Kluge, 1914), Nematoflustra flagellata (Waters, 1904) and Himantozoum antarcticum (Calvet, 1905). Approximately ten colonies of each species were collected monthly by SCUBA divers over a 14 mo period during 1992/1993, and the status of each individual zooid was classified as differentiating/regenerating, active (feeding autozooids), degenerate (brown body) or sexually reproductive (ovicells present, or zooid containing a larva). Polypide cycling in all three species was distinctly seasonal. New zooids formed at the growth margin and typically contained actively feeding polypides for ≃9 mo before these polypides degenerated into brown bodies in the austral winter (June). Very few polypides were active in the period from June to August, when water-column food levels were at their lowest; after this period new polypides differentiated. Individual zooids typically underwent a total of five (I. rubefacta and N. flagellata), or at least four (H. antarcticum) complete polypide cycles before becoming senescent. Polypide lifetimes generally became shorter as the age of the zooid increased. Sexual reproduction was also distinctly seasonal in these species, with bands of ovicells or sexually reproductive zooids being formed each year in late summer once a given colony had grown to a threshold size (or age). Larvae were then brooded for ≃10 mo before being released in January/February (N. flagellata) or February/March (H. antarcticum). The seasonal patterns of polypide cycling are related clearly to the variations in food availability, and these species appear to have the longest zooid lifetime (≃5 yr) and the slowest polypide cycling (once per year with polypide lifetimes up to 10 mo) reported for any bryozoan so far. Received: 7 May 1995 / Accepted: 19 November 1997     

Interpreting seawater temperature range using oxygen isotopes and zooid size variation in Pentapora foliacea (Bryozoa)

This is the first investigation of how two independent proxies for seawater temperature inference (zooid size variation and oxygen isotope ratios of skeletal carbonate) relate to the actual measured ranges of temperature experienced by cheilostome bryozoan colonies. Nine specimens of the bimineralic marine cheilostome bryozoan Pentapora foliacea (Ellis and Solander, 1786) were analysed, collected from ~18-m depth at two localities in Wales, UK—four from Skomer Island (51°42′510″N, 5°13′42.60″W) and five from Porth Ysgaden, Lleyn Peninsula (52°54′6.75″N, 4°38′47.34″W). The annual range of temperature implied by zooid size variability provides a good approximation of the actual range of temperature recorded by a datalogger. However, annual ranges of temperature reconstructed from skeletal oxygen isotope ratios were narrower, typically not showing the lowest temperatures experienced by the colonies. This can be explained by progressive thickening of zooid skeletal walls during the life of the colony that homogenises the temperature signal by time-averaging over the lifetime of the colonies. Our study provides evidence that a combined morphological isotope approach has great potential in the reconstruction of annual ranges in seawater temperatures from historical and fossil bryozoans, particularly for species that lack ontogenetic skeletal wall thickening and bimineralic skeletal composition. As cheilostome bryozoans have been common in benthic communities since the Late Cretaceous, they represent a valuable and underutilised resource for the interpretation of environmental regimes.     

Influence of sea surface temperature on reproductive period and size at maturity of brown shrimp (<Emphasis Type="Italic">Farfantepenaeus californiensis</Emphasis>) in the Gulf of California

Reproductive period and size at maturity of the brown shrimp Farfantepenaeus californiensis were analyzed for correspondence to sea surface temperature (SST) in three zones of the Gulf of California. Mature females from fishing areas in the north (Puerto Peñasco), center (Guaymas), and south (Mazatlán), and monthly SST were examined. Average SST for 1983–2000 decreased from Mazatlán (26.2±0.2°C) to Puerto Peñasco (22.6±0.3°C). The seasonal variation in SST between coldest and warmest months was 7.8°C in Mazatlán and 11.4°C in Puerto Peñasco. The size of shrimps at maturity was inversely correlated to SST, increasing from Mazatlán (121 mm total length) to Puerto Peñasco (154 mm total length). The reproductive period near Mazatlán is year-round. Guaymas and Puerto Peñasco have one period of high intensity. We conclude that warmer water and low seasonal variability allow brown shrimp to reproduce more frequently, but reach smaller size at maturity.Communicated by P.W. Sammarco, Chauvin     

Seasonal reproductive biology of the egg-carrying calanoid copepod Pseudodiaptomus marinus in a eutrophic inlet of the Inland Sea of Japan

In situ egg production of the egg-carrying calanoid copepod Pseudodiaptomus marinus was investigated in Fukuyama Harbor, a eutrophic inlet of the Inland Sea of Japan, at 3- to 5-d intervals for a year. This species reproduced throughout the year, and the adults showed a large abundance peak in June/July and a small peak in September/October. Females usually outnumbered males, comprising 61.4% of the annual mean. The composition of ovigerous females varied from 7.9 to 100%, with an annual mean of 55.7%. Adult prosome length was consistently large throughout winter and spring, and decreased with increasing temperature in summer and fall. Egg diameter varied from 98 to 121 μm, and was negatively correlated to temperature. The seasonal variation in clutch size (range: 15.1 to 38.2 eggs) was bicyclical, with peaks in May and December. The egg production rate of breeding females was low in January to March (mean: 2.3 eggs female⁻¹ d⁻¹), while it was constantly high from mid-May to early October (mean: 12.1 eggs female⁻¹ d⁻¹). The specific egg production rate for the breeding females was highly correlated to temperature; it increased linearly from 0.03 d⁻¹ at 9 °C to 0.27 d⁻¹ at 26 °C. Compared to other co-occurring copepods, the reproductive rate of P. marinus was lowest, which is one of the reasons why this species never dominates in this inlet. Received: 11 November 1996 / Accepted: 7 December 1996     

Otolith age validation and growth-rate variation in flyingfish (Hirundichthys affinis) from the eastern Caribbean

A study of otolith aging and growth-rate variation in the flyingfish Hirundichthys affinis (Günther) was conducted in the eastern Caribbean (10–16°N; 58–62°W) in 1987–1989. Daily otolith-increment formation was validated in laboratory-reared larvae, confirming the usefulness of otolith-increment counts for age determination of H. affinis juveniles (<150 mm fork length, FL). A mark-recapture programme to validate increment formation in wild adults was unsuccessful due to tetracycline-linked mortality and insufficient tetracycline uptake in slow-growing adult otoliths. A von Bertalanffy growth curve fitted to juvenile size-at-age data gave preliminary growth-curve parameters of t ₀=2.85 d and k=0.00854 on a daily basis, with an asymptotic length, L, of 245 mm FL, for eastern Caribbean flyingfish. Juvenile growth rate in H. affinis is sensitive to spatial and temporal variation in temperature. Growth rates were higher where sea-surface temperatures were higher, and were higher for juveniles hatched in warmer months (April–July) than in colder months (November–March). Growth rates were also higher near islands than at more oceanic locations. Variation in juvenile growth rates may influence the spatial and temporal variation in spawning frequency observed in H. affinis.     

Reproductive cycle and gametogenesis in the black abalone Haliotis cracheroidii (Gastropoda: Prosobranchiata)

Seasonal changes in gonald size and stages of gametogenesis in the black abalone Haliotis cracheroidii were related to changes in environmental parameters. H. cracheroidii showed an annual reproductive cycle terminating in a synchronized spawning in late summer. Gametogenesis was initiated immediately after spawning. Gametes were present in the gonad through the winter months. Gametogenesis was initiated a second time in the spring months. Maximal gonad growth (to a gonad index of 20%) occurred during summer months prior to spawning. Changes in gonad size andperiods of initiation of gametogenesis revealed no apparent correlation with changes in seasonal water temperature. Changes in gonad size showed no apparent relation to change in day length. Total polysaccharide levels in foot tissue changed seasonally, indicating that food availability is probably not a factor in directly regulating gonad growth. Gonad index data for the chiton Katharina tunicata (collected over a 10 year period) showed no apparent correlation to seasonal change in water temperature.     

Cephalopod hatchling growth: the effects of initial size and seasonal temperatures

Temperature is known to have a strong influence on cephalopod growth during the early exponential growth phase. Most captive growth studies have used constant temperature regimes and assumed that populations are composed of identically sized individuals at hatching, overlooking the effects of seasonal temperature variation and individual hatchling size heterogeneity. This study investigated the relative roles of initial hatchling size and simulated natural seasonal temperature regimes on the growth of 64 captive Octopus pallidus over a 4-month period. Initial weights were recorded, and daily food consumption and fortnightly growth monitored. Two temperature treatments were applied replicating local seasonal water temperatures: spring/summer (14–18°C) and summer/autumn (18–14°C). Overall octopuses in the spring/summer treatment grew at a rate of 1.42% bwd⁻¹ (% body weight per day) compared to 1.72% bwd⁻¹ in the summer/autumn treatment. Initial size influenced growth rate in the summer/autumn treatment with smaller octopuses (<0.25 g) growing faster at 1.82% bwd⁻¹ compared to larger octopuses at 1.68% bwd⁻¹. This was opposite to individuals in the spring/summer treatment where smaller octopuses grew slower at 1.29% bwd⁻¹ compared to larger octopuses at 1.60% bwd⁻¹. Initial size influenced subsequent growth, however, this was dependent on feeding rate and appears to be secondary to the effects of temperature.     

Seasonal variation in reproduction and population structure of the bivalve Limopsis tajimae in the upper bathyal zone of Suruga Bay, Japan

Seasonal variation in reproduction and population size structure was investigated for the suspension-feeding bivalve Limopsis tajimae Sowerby inhabiting the upper bathyal zone (300 m deep) of Suruga Bay, central Japan. The bivalve was collected at 1- to 4-month intervals for a period of 22 months, and bottom environment was monitored concurrently to detect factors affecting seasonality in the bivalve. Bottom water temperature, organic carbon and nitrogen contents in the sediments did not exhibit seasonal variation. Size-adjusted soft-tissue weight varied slightly, but statistically significantly between stations and months. However, its seasonal pattern was not obvious, and the pattern of temporal variation was totally different between stations. The sex ratio did not deviate from 1:1, and there was no significant difference between shell lengths of females and males. Females possess both immature small oocytes and large developed oocytes in their ovaries throughout the year, suggesting that they can potentially undergo year-round continuous reproduction. The proportion of developed oocytes in each female varied greatly from month to month, although no seasonal cycle was obvious. Population size structure of L. tajimae was polymodal. A mode of the smallest size class occurred in most months, suggesting long periods of bivalve recruitment. These findings indicate that seasonal variation in reproduction of the bivalve was negligible, probably reflecting constant physical and nutritive conditions of the bottom environment. Received: 6 January 1998 / Accepted: 19 May 1998     

Influence of food quantity and temperature on development and growth of the marine copepod Calanus chilensis from northern Chile

An experiment under laboratory conditions was conducted to test the hypothesis that development and growth of copepodite stages in Calanus chilensis are temperature-dependent and not subject to food shortage in the upwelling area of the Humboldt Current, northern Chile. Field data obtained from June 1994 to May 1995 in Bahía Mejillones (23°S) were used to define four combinations of temperature and food under which copepodites were reared from Stage CIII to adulthood. The high temperature was 18.1 °C and the low temperature 13.1 °C, whereas the high food level was in the range of 6.8 to 24.8 μg l⁻¹ chlorophyll a and the low level 1.0 to 6.8 μg l⁻¹ chlorophyll a. As food a mixture of three unknown species of phytoflagellates and the diatom Navicula cryptocephala was used. This phytoplankton was initially obtained from the same sampling sites as copepods and kept in f/2 media at stable levels and composition throughout the experiment. The development rate (1/t), estimated from the time (t) elapsing between Stage CIV and adult, was significantly affected by both temperature and food, although low-food effects were much more remarkable. Low-food conditions also significantly reduced body length and “structural” (lipid-discounted) body mass at adulthood, while temperature only affected body length. The weight-specific growth rate was also affected by food and temperature, but again food effects were much more drastic. The results indicate that C. chilensis is a highly sensitive species to lack of food, and is possibly subject to food shortage during its annual cycle in the coastal upwelling area of northern Chile. Food limitation may help explain the seasonal pattern of adult size reported by previous studies in the area and the lack of consistence between the number of generations predictable from a temperature-dependent model and that observed in the field during the annual cycle. Received: 10 September 1996 / Accepted: 29 October 1996     

Vertical distribution of Calanus finmarchicus and C. helgolandicus in relation to the development of the seasonal thermocline in the Celtic Sea

The geographical distribution and annual mean abundance of Calanus finmarchicus (Gunnerus) and C. helgolandicus (Claus) in the northern North Atlantic Ocean were shown in relation to the seasonal and annual fluctuations of abundance of the species in the Celtic Sea from 1960 to 1981. These congeneric copepods, although showing allopatric distributions over most of their geographical range, have sympatric distributions in the Celtic Sea where they dominate the dry weight biomass of the plankton throughout the year. The two species respond differently to the development of the seasonal thermocline and halocline by taking up different vertical distributions in the water column. C. finmarchicus occurred in the colder, more saline water below the thermocline, while C. helgolandicus occurred in the warmer, less saline water above the thermocline. This behaviour is postulated as a mechanism by which these morphologically similar copepods more fully exploit the resources of their temporally and spatially heterogeneous environment and also minimise interspecific competition. The species have the same foraging techniques and are able to exploit the same size spectrum of particulates. The vertical depth strata in which the populations are found for most of the year in the Celtic Sea means that both species exploit the diatom bloom in early spring but, thereafter, C. helgolandicus grazes on the daily production of the autotrophs in the euphotic zone while C. finmarchicus, below the thermocline, has to rely for its food on sedimenting particulates (whole cells, detritus and faecal material). The isolating mechanisms whereby these two populations partition the habitat in the Celtic Sea are discussed.     

Temporal and spatial variation in protein,carbohydrate, and lipid levels in <Emphasis Type="Italic">Corallium rubrum</Emphasis> (Anthozoa,Octocorallia)

Red coral (Corallium rubrum) is an important component of the Mediterranean coralligenous habitat. The present study analyzes the relationship between energy storage molecule concentration (i.e. tissue balance of protein–carbohydrate–lipid levels) and the gonadal output in two populations located at different depths (16–18 m and 40–45 m depth) through a seasonal cycle (2002–2003). In June 2003, another six populations have been sampled to compare medium-scale geographical variation. The colonies sampled were less than 6 cm high, thus representing the majority of the population in the studied area. There was no clear relationship between energy storage (i.e. carbohydrate and lipid levels) and gonadal output, nor in the seasonal cycle or the spatial variation. A clear-cut seasonal trend of energy storage was detected only in the shallower population. The water temperature was warmer at 20 m depth and the water transparency (Secchi disk) was also higher (lower seston concentration) in 2003 when the same periods in 2002 and 2003 were compared (May–August). Carbohydrates seemed to be more sensitive to food constraints than lipids. The partial mortality detected in summer 2003 in the sampling area appeared to be recorded in the carbohydrate and lipid levels of this species, especially in the shallow population. The biochemical levels prove to be a useful tool to detect unusual environmental conditions that may cause partial or total mortality in benthic suspension feeders.     

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     

Thermal effects on swimming activity and habitat choice in juvenile Pacific cod (Gadus macrocephalus)

The behavioral responses of fishes to temperature variation have received less attention than physiological responses, despite their direct implications for predator–prey dynamics in aquatic ecosystems. In this paper, we describe the temperature dependence of swimming performance and behavioral characteristics of juvenile Pacific cod (Gadus macrocephalus; 75–125 mm total length). Maximum swimming speeds increased with temperature and body size. Routine swimming speeds of Pacific cod in small groups of similarly sized fish (N = 6) increased with body size and were 34 % faster at 9 °C than at 2 °C. The response to temperature was opposite that previously described for juvenile walleye pollock (Theragra chalcogramma), reflecting species-specific differences in behavioral responses. In a separate experiment, we demonstrated the effect of temperature on habitat selection of juvenile Pacific cod: Use of an artificial eelgrass patch in a 5-m-long laboratory tank was significantly greater at 9 °C than at 2 °C. These results illustrate that temperature affects a range of behavioral traits that play important roles in determining the frequency and outcomes of predator–prey interactions.     

Can marine mammals be used to monitor oceanographic conditions?

The breeding performance of higher predators has often been used to monitor fluctuations in the abundance of important prey stocks in marine ecosystems. The development of electronic data-loggers in recent years has also provided the opportunity of using wide-ranging marine animals to measure physical oceanographic conditions. In this study, time–depth recorders (TDRs) programmed to record temperature were deployed on female Antarctic fur seals (Arctocephalus gazella) at Bird Island, South Georgia (54°00′S; 38°02′W) during the breeding seasons 1994 to 1998. Temperature sensors had relatively slow response times, and thermal radiation errors occurred during the day when seals spent a large proportion of their time at the surface. Nevertheless, measurements provided temperature–depth profiles which were typical of the vertical stratification of the ocean. During the early stages of a foraging trip temperature increased, suggesting that fur seals travelled northwards from South Georgia towards the warmer waters of the Polar Front. In addition, higher temperatures were recorded by females that remained at sea for longer, implying that these individuals also travelled further. Mean sea-surface temperature (SST) increased from ∼1 to 4 °C from December to March and agreed with SSTs from ship, buoy and satellite. Future studies on marine mammals which combine satellite tracking with oceanographic measurements are likely to provide valuable information on biophysical aspects of the ocean. Received: 16 June 1998 / Accepted: 13 February 1999     

A multi-factorial experiment on heart rate variations in the intertidal crab Pachygrapsus marmoratus

A multi-factorial experiment was designed to investigate the effect of the following factors on the cardiac activity of the intertidal crab Pachygrapsus marmoratus: respiratory medium (air, water), temperature (four levels, 10, 17.5, 25, 32.5 °C), season (winter, summer) and body size (two levels, carapace ≤21 mm and carapace >21 mm). The results showed that the heart rate of P. marmoratus increased linearly with temperature and decreased when the specimens were exposed to air rather than water. Moreover, the heart rate values in summer were lower than those in winter at the corresponding temperature and body size. The summer heart rate–temperature regression line was laterally (to the right) shifted with respect to the winter line, suggesting a seasonal acclimation. Body size affected heart rate only at the acclimation temperature (17.5 °C), while no significant effect was detected at lower or higher temperatures. During the reproductive season a separate experiment was carried out to assess the effect of sex and reproductive status on heart rate. No significant difference was found among mean values of males, berried females and females without eggs. The results of the present study confirm the high physiological plasticity of this species, suggesting that P. marmoratus is a truly amphibious crab, able to deal with both water- and air-breathing during its activity. Received: 5 January 1999 / Accepted: 7 July 1999     

Temperature acclimation and other influences on embryonic duration in the copepod Pseudocalanus sp.

Embryos of Pseudocalanus sp. captured near Halifax, Nova Scotia, Canada, in spring take longer to develop than do those from the warmer waters of summer and autumn. This is related to the larger size of females and their eggs in the colder seasons. An experiment revealed size-assortative mating and the influence of male size as well on embryonic duration. The seasonal change is contrary to the slight reduction of embryonic duration after short-term cold acclimation of females in the laboratory. There is no evidence that embryos in the cold season develop more rapidly than expected from the overall relationship between embryonic duration and female size. However, they do, like embryos after short-term cold acclimation, develop relatively more rapidly at low temperatures; i.e., there is clockwise rotation of the rate-temperature response, equivalent to the temperature adaptation that has been demonstrated among species of copepods. This seasonal rotation may also be related to seasonal variations in size, since larger females from a given season produce embryos that show such rotation. Thus, there is no evidence for seasonal compensation as a consequence of temperature acclimation underlying changes that are attributable to differences in size. Evidently there is also a strong genetical component in duration of development of embryos produced by individual females.     

Changes in the timing of otolith zone formation in North Sea cod from otolith records: an early indicator of climate-induced temperature stress?

We examine the seasonal variation in otolith increment formation in southern North Sea cod as a means of monitoring how changes in sea temperature over the past 20 years have affected cod in the wild. Seasonal opaque zone formation was related to winter and early spring. Timing of opaque zone formation was not influenced by either temperature or fish length, but increasing age led to slightly earlier but slower opaque zone formation. In contrast, there was a clear shift in the timing of translucent zone formation with temperature. In warm years, translucent growth occurs up to 22 days earlier than in colder periods. Increasing age and smaller size-at-age resulted in an earlier transition from opaque to translucent edge formation. Translucent zone formation appears indicative of increasing metabolic stress, and the earlier onset provides direct evidence of the impact of increasing sea temperatures on wild North Sea cod stocks.     

Seasonality and dynamics in coral reef macroalgae: variation in condition and susceptibility to herbivory

Seasonal variation in coral reef macroalgal size and condition is well documented, yet seasonal variability of herbivory on macroalgae by coral reef fishes is unknown. Herbivore feeding intensity was quantified monthly on an inner-shelf reef on the Great Barrier Reef, using Sargassum bioassays. Removal rates of transplants displayed high levels of variation with significantly higher rates of removal during the summer months. Differences in Sargassum plant size and condition suggest that the variability in herbivore feeding intensity is attributed primarily to the variation in the condition of the macroalgae, especially epiphyte loads. The dramatic changes in macroalgal removal reveal a considerable decrease in herbivore activity in the winter. This highlights the clear distinction between ‘summer’ and ‘winter’ months in terms of reef processes, emphasizing the high seasonal variation in macroalgal removal rates at different time of the year.