Feeding by larvae of two different developmental modes in Streblospio benedicti (Polychaeta: Spionidae)

Females of the spionid polychaete Streblospio benedicti (Webster) produce either small eggs (60–70 μm diameter) and planktotrophic larvae, or large eggs (100–200 μm) and lecithotrophic larvae that reportedly do not feed. This intraspecific polymorphism, a form of poecilogony, is potentially useful in studies of larval ecology and evolution, but necessary data on larval form and function are lacking. This study describes the morphology and nutritional biology of larvae obtained from Atlantic (South Carolina) and Pacific (California and Washington) populations from 2003 to 2005. The two types of larvae produced by Atlantic S. benedicti differed greatly in length (229±22 μm SD for planktotrophs vs. 638±40 μm for lecithotrophs) and chaetiger number (2–5 vs. 10–11) at release from the female’s brood pouch. Planktotrophic larvae bore long provisional chaetae on their first chaetiger; provisional chaetae were absent in lecithotrophic larvae. Larvae from Pacific populations were all of the lecithotrophic form, and were similar to their Atlantic counterparts in all respects. High-speed video microscopy revealed that both types of larvae used opposed bands of cilia to capture suspended particles and transport them to the mouth, where they were often ingested. Lecithotrophic larvae reared with suspended phytoplankton (Rhodomonas sp., 10⁴ cells ml⁻¹) for 2 days grew significantly faster than sibling larvae reared without added food, indicating that these larvae can digest and assimilate ingested food. Larvae of S. benedicti that develop from large eggs are thus facultative planktotrophs instead of obligately non-feeding lecithotrophs, a result that affects the interpretation of comparative studies of the ecology and evolution of larvae in S. benedicti and certain other marine invertebrates.     

Fuels for development: evolution of maternal provisioning in asterinid sea stars

For marine invertebrates, larval developmental mode is inseparably linked to the nutritional content of the egg. Within the asterinid family of sea stars there have been multiple, independent, evolutionary transitions to lecithotrophic development from the ancestral, planktotrophic state. To investigate the evolution of maternal investment and development within the Asterinidae, we quantified individual lipid classes and total protein for eggs and larval stages of closely related species representing three developmental modes (planktotrophy, planktonic lecithotrophy and benthic lecithotrophy). Within species, maternal provisioning differed between females indicating that egg quality varied with parentage. Maternal investment was related to egg size but, after correcting for egg volume, we identified two major oogenic modifications associated with the evolution of lecithotrophic development: (1) a reduction in protein deposition that probably reflects the reduced structural requirements of nonfeeding larvae, (2) an increase in deposition of a single class of energetic lipid, triglyceride (TG). The exception was Parvulastra exigua, which has benthic, lecithotrophic development and lays eggs with a lipid to protein ratio close to that of planktotrophs. This oogenic strategy may provide P. exigua larvae with a protein “weight-belt” that assists in maintaining a benthic existence. Asterinids with planktotrophic development used a significant portion of egg TG to build a feeding bipinnaria larva. For Meridiastra mortenseni, female-specific differences in egg TG were still evident at the bipinnaria stage indicating that egg quality has flow-on effects for larval fitness. In lecithotrophic asterinids, TG reserves were not depleted in development to the larval stage whereas protein stores may help fuel early larval development. Available data indicate that there may be two evolutionarily stable egg lipid profiles for free-spawning, temperate echinoderms.     

Cryptic invasion of northeast Pacific estuaries by the Asian polychaete, Hediste diadroma (Nereididae)

The estuarine polychaete, Hediste diadroma (Nereididae), of eastern Asia was found at all six sites sampled in October 2009 in northeast Pacific estuaries, including the Columbia River, along a 300-km stretch of coastline in Washington and Oregon, USA. The immature worms were returned to the laboratory in Japan where 13 of them were cultured to sexual maturity after 5–7 months. These worms had the diagnostic, epitoke-specific spinigers that distinguish H. diadroma from all other Hediste species. Their egg sizes (140–170 μm), paragnath numbers on the proboscis, and chromosome number (2n = 28) were also consistent with those previously reported for the Japanese population of H. diadroma. The widespread distribution of H. diadroma indicates that it is a successful invader of the North American Pacific and it may have long escaped detection because of its morphological similarity to the native species, H. limnicola. The long pelagic life of H. diadroma larvae could have facilitated its successful trans-ocean dispersal with ballast water of ships.     

Parthenogenetic reproduction of Diaphanosoma celebensis (Crustacea: Cladocera): influence of salinity on feeding, survival, growth and neonate production

 Effect of salinity on the feeding rate and parthenogenetic reproduction of asexual females of the cladoceran Diaphanosoma celebensis Stingelin was studied. Short-term (10 h) grazing experiments were conducted using Isochrysis galbana as feed at 5, 17, 25 and 30 psu salinity. Gut pigment concentration showed a significantly higher rate of feeding at lower salinities. Survival, growth, maturity attainment and neonate production of asexual females reared in the above four test salinities indicated preference for lower salinities (5 and 17 psu). The mean size of adult females decreased from 909 to 593 μm, mean life span from 24 to 5 d, mean neonate production from 12 to 2 and mean size of neonates from 434 to 400 μm as the salinity increased from 5 to 30 psu. Salinity variations also affected the size and age of primiparous females. Resting egg formation and sexual reproduction did not occur at the tested salinities. The results indicate that D. celebensis is adapted to low saline, estuarine environments. Received: 14 January 2000 / Accepted: 24 March 2000     

The effects of temperature and salinity on reproductive success of <Emphasis Type="Italic">Temora longicornis</Emphasis> in the Baltic Sea: a copepod coping with a tough situation

At specific locations within the Baltic Sea, thermoclines and haloclines can create rapid spatial and temporal changes in temperature (T) and salinity (S) exceeding 10°C and 9 psu with seasonal ranges in temperature exceeding 20°C. These wide ranges in abiotic factors affect the distribution and abundance of Baltic Sea copepods via species-specific, physiological-based impacts on vital rates. In this laboratory study, we characterized the influence of T and S on aspects of reproductive success and naupliar survival of a southwestern Baltic population of Temora longicornis (Copepoda: Calanoida). First, using ad libitum feeding conditions, we measured egg production (EP, no. of eggs female⁻¹ day⁻¹) at 12 different temperatures between 2.5 and 24°C, observing the highest mean EP at 16.9°C (12 eggs female⁻¹ day⁻¹). Next, the effect of S on EP and hatching success (HS, %) was quantified at 12°C for cohorts that had been acclimated to either 8, 14, 20 or 26 psu and tested at each of five salinities (8, 14, 20, 26 and 32 psu). The mean EP was highest for (and maximum EP similar among) 14, 20 and 26 psu cohorts when tested at their acclimation salinity whereas EP was lower at other salinities. For adults reared at 8 psu, a commonly encountered salinity in Baltic surface waters, EP was relatively low at all test salinities—a pattern indicative of osmotic stress. When incubated at 12°C and 15 different salinities between 0 and 34 psu, HS increased asymptotically with increasing S and was maximal (82.6–84.3%) between 24 and 26 psu. However, HS did depend upon the adult acclimation salinity. Finally, the 48-h survival of nauplii hatched and reared at 14 psu at one of six different temperatures (10, 12, 14, 16, 18 and 20°C) was measured after exposure to a novel salinity (either 7 or 20 psu). Upon exposure to 7 psu, 48-h naupliar mortality increased with increasing temperature, ranging from 26.7% at 10°C to 63.2% at 20°C. In contrast, after exposure to 20 psu, mortality was relatively low at all temperatures (1.7% at 10°C and ≤26.7% for all other temperatures). An intra-specific comparison of EP for three different T. longicornis populations revealed markedly different temperature optima and clearly demonstrated the negative impact of brackish (Baltic) salinities. Our results provide estimates of reproductive success and early survival of T. longicornis to the wide ranges of temperatures and salinities that will aid ongoing biophysical modeling examining climate impacts on this species within the Baltic Sea.     

Effects of salinity and possible interactions with temperature and pH on growth and photosynthesis of <Emphasis Type="Italic">Halophila johnsonii</Emphasis> Eiseman

The effects of salinity, temperature, and pH variations on growth, survival, and photosynthetic rates of the seagrass Halophila johnsonii Eiseman were examined. Growth and survival responses to salinity were characterized by aquarium experiments in which plants were exposed to seven different salinity treatments (0, 10, 20, 30, 40, 50, and 60 psu) during 15 days. Photosynthetic behavior was assessed for short-term salinity exposures (1 or 20 h) by incubation experiments in biological oxygen demand (BOD) bottles and by measuring photosynthesis versus irradiance (PI) responses in an oxygen electrode chamber. In the bottle experiments the possible effects of interactions between salinity and temperature (15, 25, and 35°C) or pH (5, 6, 7, and 8.2) were also examined. Growth and survival of H. johnsonii were significantly affected by salinity, with maximum rates obtained at 30 psu. Salinity also altered the parameters of the PI curves. Light-saturated photosynthesis (P _max) and the photosynthetic efficiency at subsaturating light (α) increased significantly up to an optimum of 40 psu, decreasing again at the highest salinities. Dark respiration rates and compensating irradiance (I _c) showed minimum values at 40 and 50 psu, while light-saturation point (I _k) was maximum at 30–50 psu. An interaction between salinity and temperature was not found although an increase of temperature alone produced an increase in α, P _max, respiration rates, and I _k. An interaction between salinity and pH was only found in the P _max response: P _max increased with pH=5 at 30 psu. In addition, reducing the pH increased α significantly. In the BOD bottles experiment a significant reduction in the dark respiration with decreasing pH was observed, but the opposite trend was observed in the photosynthetic rate. These results suggest that the endemic seagrass H. johnsonii could be negatively affected by hypo- or hypersalinity conditions, although salinity changes did not seem to alter the tolerance of this species to other environmental factors, such as temperature or pH.     

The effects of temperature and salinity on egg production and hatching success of Baltic Acartia tonsa (Copepoda: Calanoida): a laboratory investigation

The functional response of the aspects of reproductive success of a southwestern Baltic population of Acartia tonsa (Copepoda: Calanoida) was quantified in the laboratory using wide ranges in temperatures and salinities. Specifically, daily egg production (EP, # female⁻¹ day⁻¹) was determined for 4 or 5 days at 18 different temperatures between 5 and 34°C and the time course and success of hatching were evaluated at 10 different temperatures between 5 and 23°C. The effect of salinity (0 to 34 psu) on egg hatching success was also examined. The highest mean rates of EP were observed between 22 and 23°C (46.8–50.9 eggs female⁻¹ day⁻¹). When studied at 18 psu, hatching success of eggs increased with increasing temperature and was highest (92.2%) at 23°C. No hatching was observed for eggs incubated at low temperatures (≤12°C) that were produced by females acclimated to temperatures ≤10°C indicating a possible thermal threshold between 10.0 and 13.0°C below which only the production of diapause (or low quality) eggs exists in this population. When tested at 18°C, the hatching success of eggs incubated at 15 different salinities increased asymptotically with increasing salinity and was maximal (81.4–84.5%) between 17 and 25 psu. The high reproductive success observed over wide ranges in temperatures and salinities in this Baltic population demonstrates one of the mechanisms responsible for the cosmopolitan distribution of this species within productive, estuarine and marine habitats.     

Embryonic,larval, and post-metamorphic development of the sea urchin <Emphasis Type="Italic">Cassidulus mitis</Emphasis> (Echinoidea; Cassiduloida): an endemic brooding species from Rio de Janeiro,Brazil

Cassiduloids are currently rare irregular echinoids with a highly conserved adult morphology. Aristotle’s lantern is present only during the post-metamorphic stage, and little is known about the early development of species in this group. Cassidulus mitis produces eggs of about 375 μm in diameter, lecithotrophic larvae with four reduced arms with skeletal fenestrated rods, cilia along the body surface, and a ciliated band on arms and lobes. Offspring is brooded among the female spines from embryo to settler’s stage. The echinopluteus larval stage is reached 6 days after fertilization, and the settler’s stage is formed at the age of 17 days. Aristotle’s lantern appears around the thirteenth day of development. The lantern is well developed and functional in settlers. It remains until at least 62 days after fertilization and can be used to acquire food from the environment. The early development of C. mitis is unusual concerning features of typical lecithotrophic larvae (such as reduced arms), but retains some features of planktotrophic larvae (such as skeletal rods and a ciliated band). Regarding egg size, early development in C. mitis seems to be transitioning from facultative lecithotrophic to typical obligate lecithotrophic pattern in echinoid larval evolution.     

Phylogeography of two Atlantic squirrelfishes (Family Holocentridae): exploring links between pelagic larval duration and population connectivity

Genetic surveys of reef fishes have revealed high population connectivity within ocean basins, consistent with the assumption that pelagic larvae disperse long distances by oceanic currents. However, several recent studies have demonstrated that larval retention and self-recruitment may be higher than previously expected. To assess connectivity in tropical reef fishes, we contribute range-wide mtDNA surveys of two Atlantic squirrelfishes (family Holocentridae). The blackbar soldierfish, Myripristis jacobus, has a pelagic juvenile phase of about 58 days, compared to about 71 days (~22% longer) in the longjaw squirrelfish, Holocentrus ascensionis. If the pelagic duration is guiding dispersal ability, M. jacobus should have greater population genetic structure than H. ascensionis. In comparisons of mtDNA cytochrome b sequences from 69 M. jacobus (744 bp) and 101 H. ascensionis (769 bp), both species exhibited a large number of closely related haplotypes (h=0.781 and 0.974, π=0.003 and 0.006, respectively), indicating late Pleistocene coalescence of mtDNA lineages. Contrary to the prediction based on pelagic duration, M. jacobus has much less population structure (φ_ST=0.008, P=0.228) than H. ascensionis (φ_ST=0.091, P<0.001). Significant population partitions in H. ascensionis were observed between eastern, central and western Atlantic, and between Brazil and the Caribbean in the western Atlantic. These results, in combination with the findings from 13 codistributed species, indicate that pelagic larval duration is a poor predictor of population genetic structure in Atlantic reef fishes. A key to understanding this disparity may be the evolutionary depth among corresponding taxonomic groups of “reef fishes”, which extends back to the mid-Cretaceous and encompasses enormous diversity in ecology and life history. We should not expect a simple relationship between pelagic larval duration and genetic connectivity, among lineages that diverged 50–100 million years ago.     

Uncommon diversity in developmental mode and larval form in the genus <Emphasis Type="Italic">Macrophiothrix</Emphasis> (Echinodermata: Ophiuroidea)

Development mode in the ophiuroid genus Macrophiothrix includes an unusual diversity of planktonic larval forms and feeding types. The modes of development for seven congeners that coexist in coral reef habitats at Lizard Island, Australia were compared using larvae generated from crosses over several reproductive seasons from 1999 to 2003. Three species (Macrophiothrix koehleri Clark, Macrophiothrix longipeda Lamarck, Macrophiothrix lorioli Clark) develop from small eggs (<170 μm) into typical obligately feeding planktonic (planktotrophic) pluteus larvae with four larval arm pairs. The remaining four species develop from larger eggs (≥230 μm) into either facultatively-feeding or non-feeding (lecithotrophic) larval forms. The facultative planktotroph (Macrophiothrix rhabdota Clark) retains the ability to digest and benefit from food but does not require particulate food to complete metamorphosis. Among the lecithotrophic species, Macrophiothrix caenosa Hoggett retains the pluteus morphology with four pairs of larval arms, but is incapable of feeding, depending instead on maternal provisions for larval development. The remaining two lecithotrophs have simplified larval morphologies with only a single pair of full length (Macrophiothrix nereidina Lamarck) or highly reduced (Macrophiothrix belli Doderlein) larval arms and no functional mouth or gut. This genus includes the first example of facultative planktotrophy in ophiuroids, the first example in echinoderms of a complete pluteus morphology retained by a lecithotrophic larva, and three degrees of morphological simplification among lecithotrophic larval forms. Egg volume varies 20-fold among species and is related to variation in feeding mode, larval form, and development time, as predicted for the transition from planktotrophic to lecithotrophic development.     

Being young in a changing world: how temperature and salinity changes interactively modify the performance of larval stages of the barnacle <Emphasis Type="Italic">Amphibalanus improvisus</Emphasis>

The fate of key species, such as the barnacle Amphibalanus improvisus, in the course of global change is of particular interest since any change in their abundance and/or performance may entail community-wide effects. In the fluctuating Western Baltic, species typically experience a broad range of environmental conditions, which may preselect them to better cope with climate change. In this study, we examined the sensitivity of two crucial ontogenetic phases (naupliar, cypris) of the barnacle toward a range of temperature (12, 20, and 28°C) and salinity (5, 15, and 30 psu) combinations. Under all salinity treatments, nauplii developed faster at intermediate and high temperatures. Cyprid metamorphosis success, in contrast, was interactively impacted by temperature and salinity. Survival of nauplii decreased with increasing salinity under all temperature treatments. Highest settlement rates occurred at the intermediate temperature and salinity combination, i.e., 20°C and 15 psu. Settlement success of “naive” cyprids, i.e., when nauplii were raised in the absence of stress (20°C/15 psu), was less impacted by stressful temperature/salinity combinations than that of cyprids with a stress history. Here, settlement success was highest at 30 psu particularly at low and high temperatures. Surprisingly, larval survival was not highest under the conditions typical for the Kiel Fjord at the season of peak settlement (20°C/15 psu). The proportion of nauplii that ultimately transformed to attached juveniles was, however, highest under these “home” conditions. Overall, only particularly stressful combinations of temperature and salinity substantially reduced larval performance and development. Given more time for adaptation, the relatively smooth climate shifts predicted will probably not dramatically affect this species.     

Larval biology of six species of the genus Conus (Gastropoda: Toxoglossa) in Hawaii,USA

Among the Hawaiian species in the genus Conus, larval life histories range from long-term planktotrophy to nonpelagic lecithotrophy. Prehatching developmental time, hatching size, pelagic period and total prejuvenile developmental time are all significantly correlated with egg size, while settling size is correlated with none of these larval characteristics. Therefore, larval life history patterns in the genus Conus appear to be determined by 2 sets of selective forces: the first influencing egg size, and the second stipulating settling size. As egg sizes become larger, hatching sizes and prehatching developmental time increase while pelagic periods and total prejuvenile developmental times decrease. Settling size appears to be related to water depth, but food requirements of newly metamorphosed juveniles may also be important.     

Reproduction and larval development of Marenzelleria viridis (Polychaeta: Spionidae)

The pelagic larvae of the polychaete Marenzelleria viridis (Verrill, 1873) are newcomers among the meroplankton of the Baltic Sea, where they sometimes achieve an abundance of up to 21x10⁶ ind m^-3 near the coast, especially in late autumn and even in winter. Benthos samples were collected in the Darss-Zingst bodden chain from April 1992 to 1993 and used to ascertain the reproductive stages of individuals. Development from fertilized egg to benthic juvenile is described on the basis of field material. Details of the morphology of various pelagic stages and the young benthic worm are presented as drawings and scanning electron microscope photographs. In 1992, development of the gametes started in mid-May. The individuals reached maturity in late September after about 20 wk. The first planktonic stage was the fertilized egg. The larvae have initially one and later two pairs of black eyes and, in the 2nd segment, a ciliated pit. Gastrotrochs are present on the third and thereafter every alternate segment. Neuropodial setae develop once the 7-setiger stage is reached. Palps appear at the 10-setiger stage, and neuropodial bidentate hooks from the 10th to 11th setiger on. Metamorphosis into the juvenile benthic stage takes place at the earliest when the 15-setiger stage has been reached. The reproductive seasons of various populations of M. viridis and the developmental patterns of their larvae are discussed, and differences are compared with relevant findings for the spionids.     

Halogenated metabolites in two marine polychaetes and their planktotrophic and lecithotrophic larvae

This study investigated the occurrence and ontogenetic changes of halogenated secondary metabolites in planktotrophic and lecithotrophic larvae and adults of two common, infaunal polychaetes, Streblospio benedicti (Spionidae) and Capitella sp. I (Capitellidae), with different life-history traits. S. benedicti contains at least 11 chlorinated and brominated hydrocarbons (alkyl halides) while Capitella sp. I contains 3 brominated aromatic compounds. These halogenated metabolites are potential defense compounds benefiting both larvae and adults. We hypothesized that: (1) planktotrophic larvae contain halogenated metabolites because they are not protected by adult defenses, (2) quantitative and qualitative variation of planktotrophic larval halogenated metabolites parallels that of adults, and (3) brooded lecithotrophic larvae initiate the production of halogenated metabolites only after metamorphosis. To address these hypotheses, volatile halogenated compounds from polychaete extracts were separated using capillary gas chromatography and identified and quantified using mass spectrometry with selected ion monitoring. All four life stages (pre- and post-release larvae, new recruits, adults) of both S. benedicti and Capitella sp. I contained the halogenated metabolites previously identified from adults. This is the first report of halocompounds identified and quantified in polychaete larvae. Allocation of potential defense compounds to offspring varied as a function of species, feeding type and developmental stage. Pre-release larvae of S. benedicti with planktotrophic development contained the lowest concentration of total halogenated metabolites (1.75 ± 0.65 ng mm⁻³), post-release and new recruits contained intermediate concentrations (8.29 ± 1.72 and 4.73 ± 2.63 ng mm⁻³, respectively), and planktotrophic adults contained significantly greater amounts (28.9 ± 9.7 ng mm⁻³). This pattern of increasing concentrations with increasing stage of development suggests synthesis of metabolites during development. Lecithotrophic S. benedicti post-release larvae contained the greatest concentrations of halometabolites (71.1 ± 10.6 ng mm⁻³) of all S. benedicti life stages and developmental types examined, while the amount was significantly lower in new recruits (34.0 ± 15.4 ng mm⁻³). This pattern is consistent with a previously proposed hypothesis suggesting a strategy of reducing potential autotoxicity during developmental transitions. Pre-release lecithotrophic larvae of Capitella sp. I contained the highest concentration of total halogenated metabolites (1150 ± 681 ng mm⁻³), whereas the adults contained significantly lower total amounts (126 ± 68 ng mm⁻³). All concentrations of these haloaromatics are above those known to deter predation in previously conducted laboratory and field trials. As a means of conferring higher larval survivorship, lecithotrophic females of both species examined may be expending more energy on chemical defenses than their planktotrophic counterparts by supplying their lecithotrophic embryos with more of these compounds, their precursors, or with energy for their synthesis. This strategy appears common among marine lecithotrophic larval forms. Received: 14 July 1999 / Accepted: 20 January 2000     

The ecological significance of lipid/fatty acid synthesis in developing eggs and newly hatched larvae of Pacific cod (Gadus macrocephalus)

The lipid/fatty acid composition of marine fish eggs and larvae is linked with buoyancy regulation, but our understanding of such processes is largely restricted to species with pelagic eggs. In this study, we examined developmental changes in the lipid/fatty acids of eggs and embryos of Pacific cod (Gadus macrocephalus), a species that spawns demersal eggs along coastal shelf edges, but as larvae must make a rapid transition to the upper reaches of the water column. Adult Pacific cod were collected in the Gulf of Alaska during the spawning season and eggs of two females were artificially fertilized with sperm from three males for each female. The eggs were subsequently reared in the laboratory to determine (1) how lipids/fatty acids were catabolized during egg and larval development, and (2) whether lipid/fatty acid catabolism had measurable effects on egg/embryo density. Eggs incubated at 4°C began hatching after 3-weeks and continued to hatch over a 10-day period, during which there was a distinct shift in lipid classes (phospholipids (PL), triacyglycerols (TAG), and sterols (ST)) and essential fatty acids (EFAs: 22:6n-3 (DHA), 20:5n-3 (EPA), and 20:4n-6 (AA)). In the egg stage, total lipid content steadily decreased during the first 60% of development, but just prior to hatch we observed an unexpected 2–3-fold lipid increase (~6–9 μg individual⁻¹) and a significant drop in egg density. The increase in lipids was largely driven by PL, with evidence of long-chained fatty acid synthesis. Late-hatching larvae had progressively decreasing lipid and fatty acid reserves, suggesting a shift from lipogenesis to lipid catabolism with continued larval development. Egg density measures suggest that lipid/fatty acid composition is linked to buoyancy regulation as larvae shift from a demersal to a pelagic existence following hatch. The biochemical pathway by which Pacific cod are apparently able to synthesize EFAs is unknown, therefore representing a remarkable finding meriting further investigation.     

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

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

Variation in larval life-history traits among reef fishes across the Isthmus of Panama

We tested the hypothesis that regional differences in oceanic productivity have led to the evolution of predictable patterns of regional variation in life-history traits of pelagic larvae of tropical reef fishes. To do so we compared larval traits (egg and hatchling size, larval growth rate and duration, and size at settlement) among closely related reef fishes from the Atlantic and Pacific coasts of the Isthmus of Panama. This comparison provides a control for phylogenetic effects because those regions shared a common fauna prior to the rise of the Isthmus ˜3.5 million years ago, subsequent to which each fauna evolved independently under a very different productivity regime. We measured larval traits of 12 benthic-spawning damselfishes (Pomacentridae: Abudefduf, Chromis and Stegastes) and 13 pelagic-spawning wrasses (Labridae: Bodianus, Halichoeres and Thalassoma). These included members of each genus on each side of the Isthmus and four sets of transisthmian sister species of pomacentrids. Among the pomacentrids we found consistent transisthmian differences in hatchling size, but not in other larval traits. Essentially the reverse pattern occurred among the labrids – larval growth and duration differed consistently among congeners in the two regions, but without consistent differences in hatchling size or size at settlement. Neither relationship is predicted by the regional-productivity hypothesis. Most of the differences were quite small. Stronger phylogenetic effects on larval traits (inter- and intrageneric variation within regions) occur in both families and evidently overwhelm any effect of regional variation in productivity. Reassessment of data that takes into account such phylogenetic effects questions previous conclusions about the existence of regional differences in larval traits among damselfishes in the West Pacific and the Caribbean. Received: 19 January 2000 / Accepted: 26 September 2000     

Influence of environmental factors and female size on reproductive output in an invasive temperate marine gastropod <Emphasis Type="Italic">Rapana</Emphasis><Emphasis Type="Italic">venosa</Emphasis> (Muricidae)

Life history and reproductive strategies influence population dynamics at the inter- and intra-specific level. Environmental conditions suitable for gonad development and spawning, the reproductive range, may be a smaller portion of the broader species distribution. The only known breeding population of veined rapa whelks (Rapana venosa) in North America is in Chesapeake Bay, USA. There is considerable interest in the potential reproductive range of this non-indigenous species given the rapa whelk’s negative impacts on commercial shellfish species in both its native and introduced ranges. Weight-specific reproductive output is described for wild caught Chesapeake Bay rapa whelks maintained in flow-through mesocosms for 2 years. Measured reproductive output within and between egg capsule deposition seasons (years) in relation to water temperature, salinity, daylength, and female size is used to describe the rapa whelk’s reproductive range. Egg capsule production is influenced by seasonal and absolute water temperatures as well as seasonal daylength cycles. Egg capsule deposition by Chesapeake Bay rapa whelks begins at water temperatures of approximately 18°C and continues for 11–15 weeks. Forty to 70% of female whelks deposited egg capsules in most weeks during this season, producing 150–200 egg capsules female⁻¹ week⁻¹. Water temperatures >28°C caused reduced egg capsule production relative to temperatures of 20–25°C. Egg capsule production was positively related to seasonal changes in daylength, and two peaks of egg capsule deposition were observed in the 2001 and 2002 deposition seasons. The combination of declining daylength and higher water temperatures in late summer was associated with the cessation of egg capsule deposition. A lower average weight specific reproductive output in 130–145 mm SL rapa whelks (average 12 ± 1%) than in 90–106 mm SL rapa whelks (average 22 ± 1% of body weight) may reflect a life history that balances the physiological costs of maintaining a large body mass with the production of many planktonic larvae from multiple clutches of egg capsules per breeding season over a 10–15-year lifespan. Estimates of the cumulative day-degree requirements corresponding to the annual initiation of egg capsule deposition were 238 and 236 for 2001 and 2002, respectively. Reproductive output and day-degree requirements for Chesapeake Bay rapa whelks were similar to values calculated from previous studies of native muricids (Eupleura caudata and Urosalpinx cinerea). A latitudinal range of 30–41° (N and S) is predicted as the realized reproductive range for rapa whelk populations on the basis of the day-degree requirements for native whelks and reproductively active invasive rapa whelk populations. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Poecilogony in an estuarine opisthobranch: planktotrophy, lecithotrophy, and mixed clutches in a population of the ascoglossan Alderia modesta

A San Diego population of the opisthobranch mollusc Alderia modesta (Lovén, 1844) exhibits poecilogony, the presence of two development modes within a single species. In spring, half of the adults spawned masses containing ∼300 eggs with a mean diameter of 68 μm. After 3 d, these egg masses hatched planktotrophic veligers with a maximum shell dimension of 116 μm. The remaining adults spawned masses containing ∼30 eggs with a mean diameter of 105 μm. These egg masses hatched after 5 to 6 d, releasing lecithotrophic larvae with a maximum shell dimension of 186 μm. About 1% of field-collected adults produced mixed clutches containing a continuum of larval sizes, spanning the size extremes of planktotrophy and lecithotrophy and hatching larvae with a mean maximum shell dimension of 152 μm. Adults producing planktotrophic and lecithotrophic larvae were interfertile, and no hybrid breakdown was observed through the F3 generation. When starved, adults which previously produced only lecithotrophic larvae switched to producing planktotrophic larvae or mixed clutches with both planktotrophic and lecithotrophic larvae. Sequence-polymorphisms from a portion of the mitochondrial cytochrome c oxidase I gene support the conclusion that the two reproductive morphs represent a single species. Most of the lecithotrophic larvae and a small percentage of the larvae from mixed clutches were metamorphically competent within 3 d of hatching. A. modesta is the only molluscan species as yet known to have both planktotrophic and pelagic lecithotrophic development within a single natural population. Received: 14 August 1997 / Accepted: 11 April 1998