Mitochondrial DNA variation and population structure in the spiny lobster Panulirus argus

Adult spiny lobsters (Panulirus argus) were collected from nine locations throughout the tropical and subtropical northwest Atlantic Ocean and examined for mitochondrial DNA (mtDNA) variation. 187 different mtDNA haplotypes were observed among the 259 lobsters sampled. Haplotype diversity was calulated to be 0.986 and mean nucleotide sequence-diversity was estimated to be 1.44%; both of these values are among the highest reported values for a marine species. Analysis of molecular variance (AMOVA) and phenetic clustering both failed to reveal any evidence of genetic structure within and among populations of P. argus. The present data are consistent with high levels of gene flow among populations of P. argus resulting from an extended planktonic larval stage and strong prevailing ocean currents.     

Genetic differentiation of the California spiny lobster Panulirus interruptus (Randall, 1840) along the west coast of the Baja California Peninsula, Mexico

Identifying reproductive stocks in commercial species is relevant to fishery management strategies. We obtained muscle samples of California spiny lobster (Panulirus interruptus) from six localities along the west coast of the Baja California Peninsula and analyzed the genetic structure using mtDNA RFLPs. Our results indicated that all localities shared the same major haplotypes and showed a spatial homogeneity in the distribution of haplotype frequencies. An analysis of molecular variance indicated that only 0.84% of the genetic variability was explained by differences among localities and was not significantly different from zero. Weak divergences were found between Bahía Magdalena, the most southerly locality, in relation to other populations. Major oceanographic processes along this coast, combined with a long larval period that supports passive transport among localities, are suggested to explain the results.     

Origin and nature of spermatophoric mass of the spiny lobster Panulirus homarus

In the spiny lobster Panulirus homarus (Linnaeus) the spermatophoric mass consists of a highly convoluted tube, containing the sperm mass and a gelatinous matrix. The spermatophoric components have their origin in the proximal vas deferens and the completed spermatophoric mass is stored within the distal dilated part of the vas deferens. In the proximal vas deferens the inner glandular epithelial cells give rise to leaf-like typhlosoles which gradually diminish in size and finally disappear in the distal vas deferens. The secretions originating from the typhlosole form the outer gelatinous matrix of the spermatophoric mass. Histochemical observations reveal that the wall of the spermatophore tube consists of neutral mucopolysaccharide, whereas the sperm mass and the gelatinous matrix are rich in acidic mucopolysaccharides. Both polyacrylamide gel electrophoresis and agarose gel electrophoresis were used to separate the acidic mucopolysaccharide (AMPS) complexed with the proteins and the simple proteins. In addition, AMPS have been isolated following the method of Rahemtulla and Lovtrup (1974) and used for electrophoresis. In both the polyacrylamide gel electrophoresis and agarose gel electrophoresis, the extracted acid mucopolysaccharides give only a single fraction which, in comparison with the standard acid mucopolysaccharides, corresponds to chondroitin sulphate. The functional significance of the chondroitin sulphate and neutral mucopolysaccharides in the spermatophoric components of P. homarus is discussed in relation to their functional role in spermatophore hardening and protection of the delicate spermatozoa during their prolonged storage on the sternum of the female.     

Control and coordination of reproduction and molting in the spiny lobster Panulirus argus

Behavioral and physiological correlates of vernal reproduction and molting were investigated experimentally in laboratory-held spiny lobsters (Panulirus argus Latreille) from Florida to determine their control by photoperiod and temperature. Behavioral measures included courtship, copulation and aggression; physiological measures included the timing and frequency of ecdysis, spermatophore emplacement, oviposition, egg hatching, and setal (pleopod) and gonadal development. Behavior resembled that observed in nature, as evidenced in the expression of complete reproductive cycles. Long daylengths and warmer temperatures enhanced courtship, spawning frequencies, and female gonadal development, although gonadal recrudescence occurred in large females irrespective of photoperiod. A significant interaction effect between photoperiod and temperature indicated a need to examine joint, non-independent factor effects upon reproductive parameters. Photoperiod and temperature did not significantly influence aggression and male gonadal development. Photoperiod did not affect molting rates significantly. However, warm temperatures enhanced molting and growth such that sub-adult and young adult lobsters (both sexes) had highest molting rates, followed by large adult males, which in turn had higher rates than reproductively active females. Female setal development correlated positively with female size, and changed abruptly at ecdysis. Although setal lengths were highly variable, setae of 9 mm or more signified sexual maturity. Reproduction and molting were size-dependent: the largest adults mated or spawned, the smallest molted, and intermediate-sized ones either mated, or molted before mating. Spiny-lobster reproductive and molting patterns were influenced by photoperiod and temperature in a complex manner depending upon sex, season, size and developmental state.     

Behavioural responses of competent lobster postlarvae to odor plumes

We determined the ability of competent laboratory-reared Homarus americanus postlarvae to use odor plumes as a potential mechanism of orientation during the habitat selection process at settlement. In a Y-maze apparatus, both swimming direction (upstream or downstream) and arm selection (control or experimental) were strongly influenced by the nature of the chemical stimulus tested. In control conditions (unconditioned water in both proximal arms of the experimental device), postlarvae swam predominantly upstream (61:39) but displayed no arm preference (48:52). When submitted to a metabolite solution of adult conspecifics or macroalgae, postlarvae swam consistently upstream (73%) and showed a significant preference for the arm providing these stimuli (69%). In contrast, postlarvae submitted to cunner (Tautogolabrus adspersus) metabolites swam predominantly downstream (68:32), and those that did swim upstream significantly avoided the experimental arm (30:70). In repeated trials with the same individuals, postlarvae behaved similarly in 80% of the assays, suggesting a low intra-individual variability in behavioural responses. It is concluded that distance chemoreception may play a role in habitat selection by lobster postlarvae at settlement.     

Temporal patterns and transport processes in recruitment of spiny lobster (Panulirus argus) postlarvae to south Florida

We used time-series analysis to identify weekly and annual patterns in the supply of spiny lobster, Panulirus argus (Latreille, 1804), postlarvae to the Florida Keys, USA, over an 8 yr period. We also investigated the relationship between postlarval influx and wind forcing as a transport mechanism using the complex vector-scalar correlation analysis. Postlarval supply had a lunar phase periodicity at 4.5 wk intervals, with postlarval abundance peaking between the new moon and first-quarter lunar phases. A distinct annual cycle of postlarval supply with two peak periods was also apparent. Cross-correlation analysis between relative postlarval abundance and a 12 mo cycle showed that the annual peak occurs in spring, centered around March. With the 12 mo periodicity removed, a smaller peak at 5 mo intervals was also well defined. Wind-forcing for 7 d prior to the time of postlarval collection was marginally correlated with postlarval abundance through the entire time-series; the association was strongest during the late fall to early spring months. The analysis indicated that postlarval supply was correlated with winds from the northeast (ca. 45°), which are associated with winter atmospheric fronts. In contrast to results reported for other spiny lobster populations, these patterns suggest that recruitment of lobster postlarvae to south Florida is predictable only at a gross level and is presumably affected by the temporally inconsistent structure of regional oceanic gyres and variability in the timing of lobster spawning in the Caribbean. Received: 24 January 1997 / Accepted: 4 March 1997     

Multiple paternity and extra-group fertilizations in a natural population of California grunion (Leuresthes tenuis), a beach-spawning marine fish

Although individuals in many fish species move to shallow waters to spawn, the California grunion (Leuresthes tenuis) is almost unique in its constitutive display of synchronous full-emergence beach spawning. During a spawning event, fish ride large waves onshore to spawn on beach land, where their eggs incubate terrestrially. Here, we employ molecular markers to ascertain how this unusual reproductive behavior impacts genetic parentage. We developed and utilized four highly polymorphic microsatellite markers to assess maternal and paternal contributions in a total of 682 progeny from 17 nests of a natural population of L. tenuis. Alleles deduced to be of paternal origin in progeny were used to determine the minimum number of sires per nest and to estimate the true number of sires per nest via Bayesian analysis. We document the following: (a) no instances of multiple maternity for progeny within a nest; (b) a high frequency of nests (88%) with multiple paternity; and (c) an appreciable fraction of nests (18%) in which the estimated number of genetic sires (as many as nine) proved to be greater than the observed number of male attendants, thus implicating occasional extra-group fertilization events. From these and other observations, we also conclude that spawning behavior in grunions may involve site choice but not explicit mate choice. In addition to providing the first analysis of molecular parentage in a beach-spawning fish, we compare our findings to those reported previously for a beach-spawning arthropod, and we discuss the forces that may be maintaining this peculiar reproductive behavior.     

Life-form population responses of a marine planktonic diatom,Chaetoceros pseudocurvisetus,to oligotrophication in regionally upwelled water

Life-form population responses of a centric planktonic diatom,Chaetoceros pseudocurvisetus Mangin, were investigated in summer 1986 and 1988 in the Izu Islands, Japan, in regionally upwelled water where nutrient concentration changed from favorable to unfavorable conditions for active growth and reproduction (oligotrophication). Two types of life form were observed: vegetative cells of healthy and unhealthy looking conditions and resting spores. The observed life-form responses were experimentally evaluated along with a depletion of limiting nutrients. The algal population ceased vegetative growth and initiated resting spore formation with a disappearance of limiting nitrate, and this life-form response seemed to be triggered by the decrease of cellular nitrogen content below a certain level. Since a large amount of silicon was required for the resting spore formation, a part of vegetative cells were unable to form resting spores and formed unhealthy looking vegetative cells under insufficient concentrations of silicic acid. Percentage shares of the resting spores in the population were linearly related to the amounts of available silicic acid. Vegetative cells which did not form resting spores showed greater mortality than resting spores by attack of bacteria and protozoa; however, vegetative cells could respond quickly to possible nutrient replenishment.     

Molt cycle alterations in behavior,feeding and diel rhythms of a decapod crustacean,the spiny lobster Panulirus argus

The crustacean molt cycle manifests extensive behavioral changes in addition to physiological and integumentary modifications. The paucity of quantitative studies led us to characterize molt stage dependent alterations in rhythmic locomotor, feeding, and agonistic behavior of subadult spiny lobster Panulirus argus held grouped and solitarily in simulated natural conditions. Non-disruptively determined molt stages were defined as proportions of intermolt duration. Significant nocturnal rhythmicity persisted through the full molt cycle, though daily form varied as a function of activity level and molt stage. A distinct early evening peak signaled initiation of foraging and walking behavior. Thereafter, rhythm amplitude either declined progressively (low activity: unimodal pattern), remained high (high activity: nocturnal plateau), or exhibited a secondary early morning peak (intermediate activity: bimodal and polymodal patterns). Activity ceased at or immediately prior to sunrise. Ecdysis was predominantly nocturnal, probably reflecting species specific spatiotemporal movement patterns and social behavior. A selective advantage of nocturnal ecdysis was postulated concerning avoidance of cannibalism and diurnal predators. Locomotor activity and feeding rates were not equivalent through the molt cycle, though both peaked in stages B₂-C₁. Locomotor activity remained high in metecdysis, decreased sharply at proecdysis initiation (D₀), reaching lowest levels in D₁-D₃, then increased in D₃ through B₁. Activity dropped steeply at ecdysis, though lobsters were capable of intense and coordinated activity. Feeding decreased slowly through metecdysis after the B₂-C₁ maximum, then increased temporarily in C₄ and D₀, indicating heightened feeding motivation. This contrasts with the locomotor activity decrease at proecdysis. Food consumption declined rapidly in D₁ and D₂ and ceased at the D₂-D₃ transition. Feeding resumed in B₁ or B₂, intensifying to maximum in late B₂. Feeding remained relatively constant within stages, whereas locomotor activity varied greatly, though both correlated with metabolic needs. Grouped and solitary lobsters displayed similar patterns of foraging and walking, equivalent to those of locomotor activity and feeding of solitary individuals. Frequency of agonistic interactions (not aggression per se) remained relatively constant through the cycle, peaking in metecdysis, though the highest relative proportion occurred near ecdysis. Lobsters then were submissive and avoided physical contact with conspecifics. Clearly, locomotor activity, feeding, and social behavior of P. argus are not simply determined. Indeed, behavior is distinctly phase coordinated with varying metabolic requirements dependent on the proximity to ecdysis and ecological pressures.     

Direct estimation of natural mortality rates for littoral marine fishes using populational data from a marine reserve

 Visual censuses conducted in a marine reserve (Medas Islands) were used to estimate the natural mortality rates (M) for five common fish species in the NW Mediterranean Sea (Coris julis, Diplodus annularis, D. sargus, Serranus cabrilla and Symphodus roissali). Visual censuses of these same five species were also performed at three sites in unprotected areas of the coast where both commercial and sport fishing activity was normal. Censuses were conducted over a 3 year period. Estimates of M in the 3 years displayed scant seasonal or interannual variation, which may mean that the populations were in equilibrium during that period. The results of this study showed that the relationships between M and the growth parameters and maximum life span were unclear, and considerable caution is therefore recommended when using indirect methods of estimating M based on those parameters. For certain species the values of M were equal to or greater than the estimated total mortality in the exploited areas. The virtual absence of piscivorous predators in the unprotected area as a consequence of the high level of fishing in that area contrasts with the high abundance of such predators in the marine reserve. Since predation is the main contributor to M, estimated mortality in the unprotected areas is attributable nearly entirely to fishing. It is suggested that M may vary according to alterations taking place in conditions in the ecosystem inhabited by a species and thus that use of a value of M for a pristine population cannot be extrapolated to exploited areas. Received: 18 January 2000 / Accepted: 14 July 2000     

Moulting and reproduction of the spiny lobster Jasus edwardsii (Decapoda: Palinuridae) in northern New Zealand

Aspects of moulting and reproduction of a primarily adult population of the spiny lobster Jasus edwardsii (Hutton) (Palinuridae) were investigated in the field and at an adjacent laboratory in northeast New Zealand from 1982 to 1986. Most males moulted in October/November, ensuring they were in intermoult stage at mating. Females moulted in late April–June, 2 to 3 mo later than females from more southern latitudes. For individual females the specific timing of ecdysis and mating and the length of the interval between them were related to size. Larger females (125 to 135 mm carapace length), tended to moult earlier and wait longer (up to 38 d) before mating, but did so prior to smaller females (95 to 105 mm carapace length, CL) which moulted later, but mated within as few as 9 d. These differences are explained equally well by size-dependent growth processes and by social or chemical inhibition by larger females. Most spiny lobsters (93 to 131 mm CL) moulted at night at depths <10 m. No clear reasons for these behaviours could be found. Eggs were incubated for 101 to 116 d by females in the study population, 70% of the time recorded for more southern areas, these differences probably being related to differences in water temperature. A consequence of earlier female moulting and egg-laying, and longer egg-incubation periods with increasing latitude is that the majority of J. edwardsii larvae hatch in September–November.     

Implementation of a marine reserve has a rapid but short-lived effect on recreational angler use

Changes in human behavior are a precursor to measurable impacts of no-take marine reserves. We investigated changes in recreational fishing site selection in response to the 2005 announcement of enforcement in a marine reserve in the Gulf of California, Mexico. We used a novel data set of daily self-reported boating destinations from emergency rescue logbooks for a recreational angling community from 2000 to 2008. Because the reserve system has no experimental control, we modeled the data two ways to test for robustness to model specification. We tested for changes in human fishing behavior with regression and fit a fleet-level discrete choice model to project a. counterfactual scenario. The counterfactual is the statistically constructed ex post expectation of the human behavior we would have observed if the reserve never existed. We included month and year fixed effects in our models to account for seasonal and interannual fluctuations in fishing behavior and catch rates. We detected a decrease in reserve use compared to the counterfactual, indicating that the reserve rapidly experienced a decrease in visitation. However, the reserve's effect to reduce trips diminished with time. These results indicate that the reserve is unlikely to meet its ecological goals without institutional changes that enhance compliance. This illustrates the value of human use data to understanding the processes underlying marine reserve function. We suggest that managers should consider human use with the same frequency, rigor, and tools as they do fishery stocks. Marine reserves directly affect people, and understanding human behavioral responses to marine reserves is an important step in marine reserve management.     

Metabolic responses of the squat lobster (Pleuroncodes monodon) larvae to low oxygen concentration

Squat lobster populations found in the Humboldt Current System over the continental shelf from ~28 to 37°S release pelagic larvae in sub-surface cold (~11 °C) hypoxic waters. Larvae subsequently spread throughout the water column encountering both normoxic and hypoxic conditions. We analyzed some short- and long-term responses of Pleuroncodes monodon larval metabolism to hypoxia at 11 °C. Routine and postprandial aerobic respiration rates were lower in hypoxia than in normoxia for all zoeal stages. Zoea V oxyconformed, while megalopae oxyregulated down to very low oxygen concentrations. Throughout zoea I development, the rate of nitrogen (protein) accumulation in zoea I was lower, and C:N ratios were higher under hypoxic conditions than in normoxia. Citrate synthase (CS) and malate dehydrogenase (MDH) apparent specific activities (as indicators of aerobic and metabolic potentials, respectively) decreased and remained at the same level, respectively, throughout zoea I reared under hypoxic conditions. Anaerobic to aerobic potential (lactate dehydrogenase (LDH)/CS) was higher in organisms reared under hypoxia, and MDH/LDH potential ratios were characteristic of organisms tolerant to hypoxia. In spite of P. monodon zoea endurance and metabolic adaptations to decreasing oxygen tensions, intense hypoxia as such of their release site would affect their overall condition especially toward the end of the molt cycle. Our results indicate the importance of considering the interaction between environmental oxygen variability and recruitment success.     

Small-scale genetic structure in a marine population in relation to water circulation and egg characteristics

Until the last decade it was assumed that most marine species have pronounced gene flow over vast areas, largely because of their potential for dispersal during early life stages. However, recent genetic, modeling, and field studies have shown that marine populations may be structured at scales that are inconsistent with extensive dispersal of eggs and larvae. Such findings have stimulated the birth of new studies explaining the mechanisms that promote population structure and isolation in the oceans, in the face of high potential for dispersal. Here we study the vertical and horizontal distribution of cod (Gadus morhua) eggs in relation to small-scale circulation and water column hydrography in a coastal location of southern Norway. Previous studies conducted in this region have shown that cod populations inhabiting fjord locations, which are on average 30 km apart, are genetically differentiated, a remarkable outcome considering that Atlantic cod have pelagic egg stages and long pelagic larval duration. We document that cod eggs are found in greater abundance in shallow water layers, which on average are flowing up the fjord (away from the open ocean), and in the inner portion of the fjord, which is subject to lower current speeds compared to the outer or mouth of the fjord. Eggs were found to be neutrally buoyant at shallow depths, a trait that also favors local retention, given the local circulation. The same patterns held during two environmentally contrasting years. These results strongly suggest that population structure of Atlantic cod is favored and maintained by a balance between water circulation and egg characteristics.     

Interactions between nutritional status and long-term responses to ultraviolet-B radiation stress in a marine diatom

Influences of nutritional status on the photoinhibitory effects of ultraviolet-B radiation (UVBR: 290 to 320 nm) on the specific growth rates (_obs) and biomass of Phaeodactylum tricornutum were determined using nutrient-replete batch cultures and nutrient-limited continuous cultures. P. tricornutum cultures were exposed to UVBR doses representative of current mid-latitude and ozone-depletion intensities. Specific growth rates and biomass were inhibited from 2 to 16% by UVBR during nutrient-replete growth. However, no effect of UVBR on _obs or biomass was detectable when nutrient limitation exceeded the potential for limitation by UVBR. Thus, a competitive interaction appears to occur between macronutrient stress and UVBR stress, such that _obs and biomass will be determined by the most limiting factor. Our results suggest that measurable decreases in phytoplankton _obs and biomass from UVBR are most likely in nutrient-rich areas of the ocean, while these parameters may not be appropriate for measuring UVBR stress in regions of nutrient limitation.     

Spring phenological responses of marine and freshwater plankton to changing temperature and light conditions

Shifts in the timing and magnitude of the spring plankton bloom in response to climate change have been observed across a wide range of aquatic systems. We used meta-analysis to investigate phenological responses of marine and freshwater plankton communities in mesocosms subjected to experimental manipulations of temperature and light intensity. Systems differed with respect to the dominant mesozooplankton (copepods in seawater and daphnids in freshwater). Higher water temperatures advanced the bloom timing of most functional plankton groups in both marine and freshwater systems. In contrast to timing, responses of bloom magnitudes were more variable among taxa and systems and were influenced by light intensity and trophic interactions. Increased light levels increased the magnitude of the spring peaks of most phytoplankton taxa and of total phytoplankton biomass. Intensified size-selective grazing of copepods in warming scenarios affected phytoplankton size structure and lowered intermediate (20–200?μm)-sized phytoplankton in marine systems. In contrast, plankton peak magnitudes in freshwater systems were unaffected by temperature, but decreased at lower light intensities, suggesting that filter feeding daphnids are sensitive to changes in algal carrying capacity as mediated by light supply. Our analysis confirms the general shift toward earlier blooms at increased temperature in both marine and freshwater systems and supports predictions that effects of climate change on plankton production will vary among sites, depending on resource limitation and species composition.     

Ontogenetic and sex-specific differences in density-dependent habitat selection of a marine fish population

The spatial dynamics of species are the result of complex interactions between density-independent and density-dependent sources of variability. Disentangling these two sources of variability has challenged ecologists working in both terrestrial and aquatic ecosystems. Using a novel spatially explicit statistical model, we tested for the presence of density-independent and density-dependent habitat selection in yellowfin sole (Limanda aspera) in the eastern Bering Sea. We found specificities in the density-dependent processes operating across ontogeny and particularly with gender. Density-dependent habitat expansion occurred primarily in females, and to a lesser degree in males. These patterns were especially evident in adult stages, while juvenile stages of both sexes exhibited a mix of different dynamics. Association of yellowfin sole with substrate type also varied by sex and to a lesser degree with size, with large females distributed over a wider range of substrates than males. Moreover, yellowfin sole expanded northward as cold subsurface waters retracted in summer, suggesting high sensitivity to arctic warming. Our findings illustrate how marginal habitats can play an important role in buffering density-dependent habitat expansion, with direct implications for resource management. Our spatially explicit modeling approach is effective in evaluating density-dependent spatial dynamics, and can easily be used to test similar hypotheses from a variety of aquatic and terrestrial ecosystems.