Delayed recovery of giant kelp near its southern range limit in the North Pacific following El Niño

The northern distribution limit of giant kelp, Macrocystis pyrifera (Agardh), along the Pacific coast of North America is relatively stable near Point Año Nuevo, California, but its southern limit has varied over hundreds of kilometers along the Baja California peninsula during the past 20 years. The factors that drive this variability in the southern limit were examined by: (1) quantifying the abundances of M. pyrifera and the subsurface stipitate kelp, Eisenia arborea (Areschoug), near M. pyriferas then southern limit at Punta San Roque (27°15N; 114°42W) on ten occasions between August 1997 and June 2002 (prior to and following the 1997/1998 El Niño Southern Oscillation); (2) initiating a field manipulation at Punta San Roque in June 2000 to examine competition between these two kelp species; and (3) surveying 20 additional sites near Punta San Hipólito (27°01N; 114°00W) in June 2002 to determine whether M. pyrifera populations had recovered to their pre-1982/1983 El Niño southern range limit after ~20 years of absence. These observations and the experimental results suggest that M. pyrifera is removed from its southern limit by the high temperatures and large waves associated with El Niños, while E. arborea is able to survive and recruit in high densities and thereby delay or even prevent M. pyriferas recovery. An inverse relationship was observed between these two species over the 5-year study. The field experiment showed that at both 4 months and 1 year after areas occupied by E. arborea were cleared, substantial recruitment of M. pyrifera occurred, while recruitment was negligible in uncleared areas. This study suggests that the southern range limit of M. pyrifera is determined by increased mortality and recruitment failure following the warm temperatures and low nutrient conditions associated with El Niños, limited substrate availability, and by competition with the more persistent populations of E. arborea.Communicated by J.P. Grassle, New Brunswick     

Effect of temperature on laboratory growth,reproduction and life span of Octopus bimaculoides

Laboratory culture of 40 Octopus bimaculoides from April 1982 to August 1983 through the full life cycle at 18°C vs 23°C provided information on the growth, reproductive biology and life span of this California littoral octopus. At 18°C, the cephalopods grew from a hatchling size of 0.07 g to a mean of 619 g in 404 d; the largest individual was 872 g. Octopuses cultured at 23°C reached their highest mean weight of 597 g in 370 d; the largest individual grown at this temperature was 848 g after 404 d. Growth data revealed a two-phase growth pattern: a 5 mo exponential phase followed by a slower logarithmic (power function) phase until spawning. At 5 mo octopuses grown at 23°C were over three times larger than their 18°C siblings. However, beyond 6.5 mo, growth rates were no higher at 23°C than at 18°C. At 13.5 mo, the mean weight of the 18°C group surpassed that of the 23°C group. The slope of the length/weight (L/W) relationship was significantly different for the two temperature regimes, with the 23°C octopuses weighing 18% less than their 18°C siblings at a mantle length of 100 mm. Females weighed more than males at any given mantle length. Males grew slightly larger and matured before females. The L/W relationship indicated isometric body growth throughout the life cycle. Higher temperature accelerated all aspects of reproductive biology and shortened life span by as much as 20% (from approximately 16 to 13 mo). O. bimaculoides has one of the longest life cycles among species with large eggs and benthic hatchlings. Extrapolations to field growth are made, and the possible effects of temperature anomalies such as El Niño are discussed.     

Reef coral reproduction in the eastern Pacific: Costa Rica,Panamá, and Galápagos Islands (Ecuador). III. Agariciidae (Pavona gigantea and Gardineroseris planulata)

The reproductive ecology of Pavona gigantea Verrill and Gardineroseris planulata (Dana) was investigated in the equatorial eastern Pacific region from 1985 to 1994. These zooxanthellate scleractinian corals were adversely affected in this region during the 1982–1983 El Niño warming event. Both species were hermaphroditic, with individual colonies showing sequential cosexual development, thus resulting in dominantly outbreeding reproduction. Sexuality was mixed, with high percentages of gonochoric and hermaphroditic colonies in both species. Approximately 1:1 male-to-female gonad ratios were found in gonochoric and hermaphroditic colonies combined. Broadcast spawning was observed in P. gigantea in the Galápagos Islands, and the sudden disappearance of mature gametes and the presence of spent gonads suggest that G. planulata is also a broadcast spawner. Colonies of both species with 200 cm² (10 cm diam) live tissue were nonreproductive. Estimated ages of the youngest reproductive colonies were 11 yr for P. gigantea and 20 yr for G. planulata. The percentage of all colonies of P. gigantea with gonads at nonupwelling sites (Caño Istand, Costa Rica and Uva Island, Panamá) ranged from 37 to 47%, respectively; colonies with gonads from upwelling environments (Saboga and Taboga Islands, Panamá) ranged from 31 to 39%, respectively, and reproductively active colonies from the thermally variable Galápagos islands comprised 40% of the collections. Compared with P. gigantea, the numbers of sexually active G. planulata colonies were roughly onehalf at nonupwelling Caño Island (20%) and Uva Island (25%) sites, or less (10%) at the upwelling Saboga Island site. Peak reproductive activity in P. gigantea occurred during the rainy season at all study sites. In the nonupwelling Costa Rican (Caño Island) and Panamainan (Uva Island) sites, mean monthly sea-surface temperatures (SSTs) were high (28 to 29°C), but slightly lower than in the dry season (29°C). In the upwelling Gulf of Panamá (Saboga and Taboga Islands), reproduction occurred after mean monthly SSTs increased from 24 to 28–29°C. In the Galápagos Islands, reproductive activity peaked during sea warming, when mean monthly SSTs reached 25°C. Sexually active colonies of G. planulata, present only at the main collection sites of Caño and Uva Islands, were also observed during the wet season. The presence of mature or spawned gonalds in both species mostly around new and full lunar phases suggests that spawning is at least weakly synchronized with moon phase. Fecundity estimates disclosed the following nonsignificant differences between sites for P. gigantea, expressed as egg production cm^-2 colony surface surface yr^-1: Galápagos (10 300 to 30 800), Uva Island (4900 to 9800), Caño Island (1800 to 7400), Saboga Island (600 to 1300) Taboga Island (1200 to 2400). Fecundity estimates for G. planulata were considerably lower: Uva Island (700 to 1400), Caño Island (500 to 1000). The sexual recruitment of P. gigantea into El Niño-Southern Oscillation (ENSO) 1982–1983-disturbed, equatorial eastern Pacific coral communities has been low, with only moderate recovery evident since 1983. G. planulata has revealed no sexual recruitment where seed populations are absent or rare (Caño Island, Galápagos Islands), and only low recruitment (Panamá) in areas with colonies that survived the ENSO disturbance.     

Nutrient-limited growth of juvenile kelp,Macrocystis pyrifera,during the 1982–1984 “El Niño” in southern California

The relative growth rates of juvenile Macrocystis pyrifera in southern California kelp forests were substantially reduced during the El Niño of 1982–1984. The lower growth rates were correlated with increased temperature and decreased nitrogen availability. Fertilization of juvenile plants with slow-release nitrogen-phosphorus fertilizer increased their growth rates to levels previously observed when temperatures were low and nutrient levels were high. The limitation in growth of M. pyrifera by levels of available nutrients during El Niño was in contrast to the usual limitation of growth by irradiance during non-El Niño years. Thus, there was a shift in the relative importance of factors controlling growth of juvenile M. pyrifera during El Niño.     

Tolerance to high temperatures and potential impact of sea warming on reef fishes of Gorgona Island (tropical eastern Pacific)

Knowledge of upper thermal-tolerance limits of marine organisms in the tropical eastern Pacific (TEP) is important because of the influence of phenomena such as El Niño and global warming, which increase sea temperature. Laboratory experiments were conducted to determine the critical thermal maximum (CTM) of reef fishes from the TEP. In 15 reef fishes of Gorgona Island (TEP) the CTM was between 34.7°C and 40.8°C. None of these CTMs was exceeded by sea temperature in the TEP during any of the strongest El Niño events in this century (32°C during El Niño 1982-1983 and 1997-1998), which indicates that all species studied here may tolerate El Niño maximum temperatures. In addition, the CTM of the least-tolerant species was 8°C above the current mean sea temperature in a wide range of latitudes in the TEP. This suggests that fishes live far from their upper thermal tolerance limits and that the current global-warming trend is still unlikely to be dangerous for these species. If sea temperature continues to increase at the current rate, in about a century sea temperature could exceed the thermal tolerance of some reef fishes and threaten them with extinction. Such risk, however, might occur sooner if the sea temperature during El Niño also increased in step with the global warming, but also because other processes involved in maintaining population, such as reproduction, can be affected at lower temperatures. The possible ability of reef fishes to adapt to increases in sea temperature is discussed.     

Inter-annual variability in somatic growth rates and mortality of coastal fishes off central Chile: an ENSO driven process?

The effects of El Niño (EN) and La Niña (LN) events upon marine organisms inhabiting the Eastern Pacific coast have been widely studied in recent years, concentrating primarily on changes in species composition and on population size. In this study, using somatic growth rates as metabolic response variables, we evaluated the mortality rates of coastal fishes inhabiting a central Chilean upwelling marine ecosystem in the South Pacific between 1990 and 2003. Four coastal fish species belonging to different trophic levels (one herbivore, one omnivore, and two carnivores) were analyzed. In all species, the estimated cohort somatic growth rates were low for those recruited during EN and high for those recruited during LN events. Annual cohort mortality rates were highest during EN events and lower during LN and transitional years. We propose that productivity (as a bottom-up driver) acts as a primary exogenous factor upon annual cohort mortality rates. We also propose that a plausible mechanism underlying this process is the negative effect the low somatic growth rates may have on fish ecological attributes such as their competitive abilities, condition factors, and predation risks, which ultimately may affect their fitness.     

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     

On the near-surface plankton of the eastern South Pacific Ocean

A study has been made of the plankton of the near-surface water layer (0 to 30 cm) of the eastern South Pacific Ocean in the region lying to the east of the meridian 90°W, between latitudes 5°N and 35°S. This region is influenced by the Peru Current: the current brings water from high latitudes, which results in a decrease in the number of species of the local fauna of copepods of the family Pontellidae, typical of tropical near-surface plankton. Some of the widely tropical and one bicentral species are absent or rare. Least affected by the Peru Current are the waters of equatorial structure in the northern part of the region. Here, 7 species of pontellids were recorded: the widely tropical Labidocera detruncata, Pontella tenuiremis, Pontellopsis regalis, the distant-neritic Pontella danae and Labidocera acuta, the bicentral Labidocera acutifrons, and the neritic Pontellopsis lubbockii. The dominant species among these are L. detruncata and L. acuta. To the west of the convergence, in the southern part of the region, live the southern central species Pontella valida and P. whiteleggei, with Pontellopsis regalis occurring occasionally. In these regions the copepod fauna is frequently dominated by pontellids. To the south of the boundary of the waters of equatorial structure, between the coast of South America and the line of convergence, lies a region most subjected to the effect of waters from high latitudes and of upwellings. It is inhabited by 2 pontellids only: Pontellopsis regalis and Labidocera acutifrons, but they too disappear close to the coast. In this particular region the copepods Calanus australis and Centropages brachiatus are common; they are found in a thicker water layer (0 to 200 m), and are often more abundant than the pontellids.     

Diet of the Humboldt penguin (<Emphasis Type="Italic">Spheniscus humboldti</Emphasis>) in northern and southern Chile

The diet of the Humboldt penguin (Spheniscus humboldti) was examined and compared in two colonies in Chile. Field work was conducted on Pan de Azúcar Island in northern Chile in the breeding season 1998/1999 and on the Puñihuil Islands in southern Chile over two successive breeding seasons during 1997/1998 and 1998/1999. Penguin diet was studied by stomach-pumping birds and analysed by species composition, size and mass of prey. Fish were the dominant prey item at both sites, the contribution of cephalopods and crustaceans varying between sites. The fish prey consisted predominantly of school fish, but there were clear latitudinal differences in fish prey taken. Penguins in the northern colony consumed primarily garfish (Scomberesox saurus), while birds at the southern colony of Puñihuil fed primarily on anchovy (Engraulis ringens), Araucanian herring (Strangomera bentincki) and silverside (Odontesthes regia). The results showed significant differences in terms of numbers of fish taken between the two breeding seasons at Puñihuil. In 1997/1998 penguins consumed almost exclusively anchovy, while they fed primarily on silversides in the successive year. Almost all prey, except stomatopods, were characterised as being pelagic species that occur in relatively inshore water, consistent with the foraging behaviour of Humboldt penguins. The dependence of Humboldt penguins on commercially exploited, schooling prey species makes the species particularly susceptible to changes in prey stocks, due to non-sustainable fisheries management.Communicated by O. Kinne, Oldendorf/Luhe     

Larval development of the red crab Pleuroncodes monodon (Decapoda Anomura: Galatheidae) under laboratory conditions

Larvae of Pleuroncodes monodon (Milne-Edwards, 1837), a red crab of commercial importance in South America, were reared in the laboratory at 2 different temperatures (15° and 20°C), from hatching up to the last larval stage. The 5 typical stages, with their corresponding functional appendages, are described and figured. The main characteristics useful in differentiating larvae of P. monodon from those of the other Chilean species of Galatheidae and its northern congener P. planipes are discussed. Data on duration of each larval stage, length of moulting intervals and mortality at the 2 test temperatures are also given.This study was financially supported by the Chilean Ministry of Agriculture and by the National Commission for Scientific and Technological Research (CONICYT).     

Annual life cycle of the copepod<Emphasis Type="Italic"> Eucalanus inermis</Emphasis> at a coastal upwelling site off Mejillones (23°S), northern Chile

Eucalanus inermis is an abundant species in the eastern tropical and subtropical South Pacific, including the oceanic and coastal waters off Chile and Peru. Its annual life cycle was studied through a time-series sampling (weekly intervals) during 2002, at a fixed coastal station at an upwelling site (Mejillones Bay, 23°S) off northern Chile. The more-or-less continuous occurrence and abundance of naupliar and copepodid stages indicated that the species reproduces during most of the year, with two peaks: one during the austral spring and the other during the summer. Thereafter, an abrupt decline in the population was observed during winter. The abundances of E. inermis copepodids and nauplii were positively correlated with sea surface temperature, suggesting temperature-dependent development and growth during the spring–summer period. Three cohorts could be distinguished during one annual cycle, with generation times >30 days. The estimate of mean weight-specific daily growth (0.12 day^–1) is lower than that of other species in the area, but it is consistent with a slower development rate. The distribution of this copepod is associated with the equatorial subsurface waters characterized by low-oxygen content (<0.5 ml l^–1). The ascent of this water mass to the near-surface during coastal upwelling in the spring–summer period and the presence of E. inermis adults favor their reproduction and the development of cohorts in the food-rich upwelling environment. The wintertime disappearance of E. inermis from shallow waters is, thus, interpreted as a movement to deep waters (>200 m depth), probably in a lethargic mode, within the oxygen minimum zone in the adjacent oceanic area.Communicated by P.W. Sammarco, Chauvin     

The effect of water movement,temperature and salinity on abundance and reproductive patterns of<Emphasis Type="Italic"> Macrocystis</Emphasis> spp. (Phaeophyta) at different latitudes in Chile

This study describes the density variation and phenology of Macrocystis integrifolia and M. pyrifera populations from northern and southern Chile, respectively. Samples of both species were taken in wave-exposed and wave-protected areas. In addition, spore production, germination and early growth rate of sporophytes of each population was studied at monthly intervals under three temperature and salinity regimes. Results indicate that M. integrifolia from northern Chile presents perennial plants with a mean density of three individuals per 0.25 m² throughout the year and that it reproduces mainly during spring and winter. Although, M. pyrifera in exposed areas of southern Chile also have a perennial-type life strategy, they are able to reproduce all year round. In contrast, M. pyrifera populations in protected areas of southern Chile show a clear annual cycle, with high recruitment during late winter and fertile sporophytes in summer and autumn, although the populations become completely decimated thereafter. The effect of temperature and salinity on M. integrifolia shows that it is independent of water movement, but requires low temperatures and high salinities for the release of zoospores, germination and early sporophyte growth. This pattern differs from that of M. pyrifera in southern Chile, which has a broader tolerance range for salinity and temperature than does M. integrifolia. However, in southern Chile wave-protected populations showed higher spore release and germination at 15°C and 18°C, whereas sporophyte growth responded better at the lowest temperature tested (8°C). In general, these results are contrary to those expected, since a seasonal reproductive pattern was observed in M. integrifolia inhabiting a less seasonally variable environment. In exposed sites of southern Chile, plants showed greater tolerance and continuous reproduction throughout the year, despite the greater environmental variability. Finally, population dynamics of protected kelps in southern Chile shows an annual pattern, which is contrary to the expected perennial strategy shown by exposed populations.Communicated by P.W. Sammarco, Chauvin     

Spatial patterns and temporal succession in soft-bottom macroinvertebrate assemblages surrounding an ocean outfall on the southern San Diego shelf: relation to anthropogenic and natural events

Spatial patterns and temporal succession in soft-bottom macroinvertebrate assemblages are described for an area of the southern San Diego shelf which is influenced by both anthropogenic and natural events. The study transect covered 19 km along the 60 m-depth contour and spanned the terminus of the Point Loma wastewater outfall (32°40N; 117°17W). Ordination and classification analyses revealed that eight major benthic assemblages occurred in the study area from 1986–1990. These analyses further indicated that: (1) the impact of wastewater discharge on the benthos was asymmetric about the outfall, and (2) primary and secondary impact zones existed within the outfall footprint. The primary impact extended only 0.5 km south, but usually up to 1 km north, of the point of discharge, reflecting the prevailing northerly flow of currents along the San Diego coast. Stations within the primary impact zone usually supported similar faunal assemblages at any point in time. A lessened, or secondary, impact was apparent at stations 1 to 2 km south and 2 km north of the outfall. The two stations 9 km to the north appeared to be outside the outfall footprint. Assemblages in the three zones did not differ greatly in species composition, but did differ with respect to organismal density, relative abundances (dominance hierarchy) of component species, and temporal persistence. Biostimulation of the benthos, manifest as increased species richness and organismal abundance, was apparent in the primary impact zone from 1986–1988, although dominance was relatively low. Dominance increased at these sites during 1989 and 1990 due to large increases in populations of a few small species, such as the polychaete Myriochele sp. M and the ostracod Euphilomedes carcharodonta. Biomass was slightly elevated in the primary impact zone during 1986–1987; however, this trend was not sustained. Communities within the outfall footprint were much less persistent over time than those beyond it. The observed shifts in benthic assemblages within the outfall footprint corresponded to natural oceanographic cycles (i.e., the transition from El Niño to La Niña conditions) and anthropogenic changes (i.e., changes in wastewater output). These observations suggest that the Point Loma outfall has a localized effect on the stability of infaunal communities. Possible destabilization mechanisms are explored. Finally, comparisons with pre-discharge data taken during 1956–1957 revealed that about half of the species present prior to construction of the outfall were still common in 1990, after 27 yr of wastewater discharge, although large increases in species diversity and abundance seem to have occurred at all stations.     

Reef coral reproduction in the eastern Pacific: Costa Rica,Panama, and Galapagos Islands (Ecuador)

Reproductive ecology of two major reef-building corals in the eastern Pacific [Pocillopora damicornis (Linnaeus) andPocillopora elegans Dana] was investigated between 1984 and 1990 in Costa Rica, Panama (Gulf of Chiriqui and Gulf of Panama), and the Galapagos Islands (Ecuador) following the 1982–83 El Niño disturbance. Mature spermatocytes and oocytes were found in both species and were usually present in the same polyp in nearly equal ratios. Numerous zooxanthellae were observed in mature, unfertilized oocytes. Although spawning in field populations was not observed, it is likely that both pocilloporids are simultaneous hermaphroditic spawners, as evidenced by the disappearance of mature gametes after full moon. This is in contrast with most known pocilloporid corals that brood and release planula larvae. Corals were reproductively most active in the thermally stable environments of Costa Rica and the Gulf of Chiriqui (Panama) where 32 to 90% of all colonies contained gametes. In the moderately varying thermal conditions in the Galapagos Islands, 16 to 40% of colonies contained gametes, and in the pronounced seasonal upwelling environment of the Pearl Islands (Panama) only 6 to 18% of colonies contained gametes. Year-round reproduction occurred in Costa Rica and the Gulf of Chiriqui, whereas reproduction was confined to warm periods in the seasonally varying environments of the Galapagos Islands and the Gulf of Panama. Pocilloporid corals in Costa Rica and the Gulf of Chiriqui demonstrated lunar spawning activity, with mature gametes present a few days before and after full moon. Some limited spawning may have occurred also at new moon. While frequent gamete maturation has been demonstrated in this study, the relatively low rates of larval recruitment occurring on eastern Pacific coral reefs disturbed by the 1982–83 El Niño suggest that the recovery of important frame-building corals could be greatly prolonged.     

A brief survey of the ecology and biology of the Caspian Polyphemoidea

Not less than 25 autochthonous species of Polyphemoidea live in the Caspian Sea; 5 of these species also inhabit the Azov and Black Seas and 3 the Aral Sea, but none is found beyond the Pontoaralocaspian basin. A great degree of polymorphism and morphological variability characterizes this group. Most Caspian Polyphemoidea exist in salinities of 12 to 13, and cannot tolerate great changes in salinity; however, 3 or 4 species in the Pontoasov basin can live in quite fresh water and populate the river reservoirs; these species do not tolerate ocean salinities over 8 to 10. All species inhabit mainly the upper layers of the sea (0 to 50 m; Cercopagis and Polyphemus exiguus down to 75 to 100 m), but avoid shallows under 5 to 15 m depth. Polyphemoidea perform diurnal vertical migrations, accumulating in surface layers during the hours of darkness and descending at sunrise; they also descend during rough weather conditions. Abundance of Polyphemoidea is subject to great seasonal variation. Most species appear in spring when the water temperature has reached 10° to 15°C; maximum abundance occurs in summer at water temperatures of 5° to 20°C, and Polyphemoidea disappear in autumn from the whole Caspian Sea except for deep areas of the South Caspian Sea, where the temperature does not drop under 10°C. Reproduction of the Caspian Podonidae is distinguished by a strikingly high rate of parthenogenesis, which is accompanied by neoteny, i.e. the embryos mature before birth. Bisexual reproduction, on the other hand, is suppressed; males and gamogenetic females containing winter eggs do not occur in all species of Podonidae and only in some species of Cercopagidae, in these latter mainly as single specimens. The majority of Cercopagidae have no males, being completely acyclic. Parthenogenesis shows a clear diurnal rhythm; delivery of young begins only after midnight and ends before sunrise. All Polyphemoidea are predators; they catch mobile prey and suck out its contents; this prevents successful observation of details of their feeding habit. However, it has been proved that Evadne anonyx feed mainly on copepods (Eurytemora) and small podonids, and this is probably true also of Cercopagis, as both these species are more marked predators than Podonidae from the open seas (Evadne nordmanni and other forms). Many aspects of taxonomy (intraspecific forms) and biology (reproduction of acyclic species, feeding, behaviour and functional morphology) are obscure and require further investigation.     

Rearing Pandalus borealis larvae in the laboratory

Larvae of the northern shrimp Pandalus borealis (Krøyer) are pelagic. In the Estuary and Gulf of St. Lawrence, Canada, the early stages are found in the upper 25-m of the water column in spring and early summer and are expected to experience a range of water temperatures from as low as 0°C to as high at 6°C. Little is known of the impact of water temperature on metabolic requirements of northern shrimp larvae. In this study, routine respiration (VO₂), maximum respiration (electron transport system activity, ETSA) and metabolic scope for growth (MS, ETSA–VO₂) of northern shrimp larvae were measured as a function of temperature (3, 5 and 8°C), developmental stage (I–V at 3°C, I–VII at 5°C and 8°C) and growth rate in dry mass. After logarithmic transformation, all three metabolic variables were linearly related to dry mass. The increase in VO₂ with body mass was faster at 5°C than at 3 or 8°C, whereas with ETSA this increase was slower. As a result, MS increased more slowly with dry mass at 5°C than at 3 and 8°C. However, MS did not limit growth in this study, since it explained only 39% of the variability in growth. All three metabolic variables as well as growth varied together as a function of temperature and ontogeny. Q₁₀ of all three metabolic variables ranged from 1.6 and 2.2 for stages I–V larvae, except for VO₂ at stage I (3.9) and stage III (2.9).     

Effects of temperature on activity,food consumption rates,and gut passage times of seaweed-eating<Emphasis Type="Italic"> Tegula</Emphasis> species (Trochidae) from California

Previous feeding studies on herbivorous marine snails rarely have focused on temperature effects on food intake. If temperature affects food intake, ectothermic snails may experience difficulty obtaining sufficient nutritional resources, limiting their ability to sustain populations at suboptimal temperatures. We hypothesized that the feeding responses of Tegula species would correspond with temperatures characteristic of their geographic distributions. We determined activity, consumption rates, and gut passage times at 11°C, 15°C, 19°C, and 23°C for three Tegula species with distinct thermal distributions: T. brunnea (cold water), T. aureotincta (warm water), and individuals from warm- and cold-water populations of T. funebralis, a broadly distributed species. Activity and consumption rates of T. aureotincta increased with increasing temperature, but were highest for T. brunnea at 19°C, a temperature rarely achieved in habitats occupied by this species, and lowest at 11°C. Warm-water T. funebralis showed significantly lower activity and consumption rates at 11°C, whereas cold-water T. funebralis consumed food fastest at 15°C and were most active at 23°C. Temperature affected gut passage time only in T. aureotincta. These data suggest that temperature might influence the northern limit of T. aureotincta by affecting activity and food consumption rates. T. brunneas activity and ability to consume food were not hindered by warmer temperatures despite the present day restriction of this species to colder waters. Also, widely separated (>300 km) T. funebralis populations may be adapted to regional conditions based on the different temperature responses of northern and southern snails.Communicated by P.W. Sammarco, Chauvin     

Reef coral reproduction in the eastern Pacific: Costa Rica,Panamá, and Galápagos Islands (Ecuador). II. Poritidae

A comparative study of the reproductive ecology of the zooxanthellate, scleractinian corals Porites lobata Dana and P. panamensis Verrill was conducted from 1985 to 1991 in eastern Pacific reef environments that were severly impacted by the 1982–1983 El Niño warming events. P. lobata, a presumed broadcast spawner of large colony size, is widely distributed in the equatorial eastern Pacific, whereas P. panamensis, a brooder of small colony size, is abundant only on some reefs in Panamá. Both species were gonochoric with nearly 1:1 sex ratios in large study populations except for P. lobata at Caño Island that had 14% hermaphroditic colonies. Mature, unfertilized oocytes contained numerous zooxanthellae in both Porites species, and all planula developmental stages contained zooxanthellae in P. panamensis. Year-round sampling revealed high proportions of colonies with gonads, ranging from 30 to 68% in P. lobata and from 60 to 68% in P. panamensis. No clear relationship between numbers of reproductive colonies and the thermal stability of the habitat was evident in P. lobata: percent colonies with gonads at non-upwelling sites was 48 to 68% at Caño Island (Costa Rica) and Uva Island (Panamá), and at upwelling sites 30 to 50% at Saboga Island and Taboga Island (Panamá), and the Galápagos Islands (Ecuador). Similarly, 90% of all P. panamensis colonies were reproductive at Uva Island (a non-upwelling site), and 86% were reproductive at Taboga Island (an upwelling site). Upwelling at Taboga Island is seasonal, nevertheless P. panamensis produced mature gonads or planulae over most of the year (11 mo), whereas P. lobata exhibited reproductive activity during only 2 mo (May and June). No clear lunar periodicity was observed in P. panamensis (Taboga Island), but a high proportion of P. lobata showed increased gonadal development around full and new moon, especially at Caño and Uva Islands. Estimated fecundities were relatively high for P. lobata at Caño (4000 eggs cm^-2 yr^-1) and Uva (5200 eggs cm^-2 yr^-1) Islands, and notably low (70 to 110 eggs cm^-2 yr^-1) in the Galápagos Islands. P. panamensis mean fecundity at Taboga Island was 720 planulae cm^-2 yr^-1 or 4.0 mm³ cm^-2 yr^-1, which was lower than the egg volume production of P. lobata at Caño and Uva Islands (7.0 to 10.0 mm³ cm^-2 yr^-1). The capacity of P. lobata and P. panamensis to reproduce sexually supports the notion that eastern Pacific coral reef recovery may not be dependent on long-distance dispersal from central Pacific areas. However, sexual recruits of P. lobata are absent or uncommon at all eastern Pacific study sites while recruits of P. panamensis were common to abundant only at the Uva Island study site. Asexual fragmentation in P. lobata augments recruitment locally, but plays no role in P. panamensis recruitment.     

Population structure and reproductive biology of Ceratonereis limnetica (Polychaeta: Nereididae) at Lower Portland,Hawkesbury River,Australia

Ceratonereis limnetica Hutchings and Glasby was collected from Lower Portland, Hawkesbury River, New South Wales, Australia, from March 1982 to September 1983. C. limnetica is one of a relatively few polychaete species in which the adults normally tolerate reverine conditions. Reproduction in this species has been modified accordingly, differing from the common pelagic spawning of many nereidids. Adults of both sexes remain essentially unmodified and spawn on the bottom. Embryos and larvae are brooded in specially constructed tubes in the sediment by the female. Both sexes have a life span of about 1 yr, dying soon after spawning. The morphological and reproductive behavioural similarities between C. limnetica and C. aequisetis are considered to indicate a close phylogenetic affinity.     

Salinity and temperature tolerance of zoeae of the portunid crab Scylla serrata

The tolerances of the first zoeal stage of the crab Scylla serrata (Forskal) have been investigated in 64 different temperature-salinity combinations. Exposure to temperatures above 25°C or to salinities below 17.5 caused considerable mortality; therefore, zoeae are unsuited to estuarine conditions. The larvae can tolerate temperatures down to 5°C is they are inactive below 10°C. It is suggested that 10°C is probably a lower limit and that female crabs which migrate to sea to release their eggs do not enter water with a temperature below 12°C. Hydrological conditions along the south-east coast of Africa indicate that females would, therefore, migrate less than 10 km offshore.