Comparative population genetic structure of marine gastropods (Littorina spp.) with and without pelagic larval dispersal

 Population genetic theory predicts that marine animal species with planktonic larvae will have less genetic structure than those with direct development. We compared the genetic structure of four species of littorinid snails – two with planktonic egg capsules that hatch as planktonic larvae and two with benthic egg masses that hatch as crawl-away juveniles. We used DNA sequencing and single stranded conformational polymorphism (SSCP) to assess sequence variation in a 480 bp fragment of the mitochondrial cytochrome b gene and then used an analysis of molecular variance (AMOVA) to estimate Φ_st for populations from the northeastern Pacific coast. One of the two direct-developing species, Littorina subrotundata, had a moderate amount of population structure (Φ_st=0.209) as expected but the other direct-developing species, L. sitkana, was nearly fixed for a single haplotype that made it impossible to precisely estimate Φ_st. One of the two planktonic-developing species, L. scutulata, did not show any significant population structure (Φ_st=0.004). In contrast to our expectations, the other planktonic-developing species, L. plena, showed some weak but statistically significant population structure (Φ_st=0.052). We discuss how differences in population genetic structure between species with the same type of development may reflect differences in their historical demography. Received: 22 December 1999 / Accepted: 24 July 2000     

Distribution pattern, reproductive traits, and molecular analysis of two coexisting vermetid gastropods of the genus Petaloconchus: a Caribbean endemic and a potential invasive species

Vermetid gastropods are characterized by complex taxonomy and unusual ecology. A survey of the fouling community in the intake channel at the Planta Centro Power Plan in Puerto Cabello, Venezuela, found massive colonies of vermetid gastropods of the genus Petaloconchus. We arbitrarily named two prevalent varieties as black and brown-orange morphs, distinguishing based on the color of their soft bodies. Spatial distribution was different for the morphs. The black morph was present along the jetty, with higher average densities in the shallower intertidal area (410 vs. 143 ind m⁻²), while the brown-orange morph was only present at the initial part of the channel (μ = 83 ind m⁻²). Both produced small eggs (142 vs. 180 μm diameter on average), with the orange-brown eggs being slightly larger, but the morphs differed in other reproductive aspects. The brown-orange morph produced significantly fewer capsules (up to nine simultaneously per female) with fewer embryos (average of 27 viable embryos/capsule) that hatched at a larger size (577 μm on average), allocating about 17.64% of the initial egg production to nurse eggs. In contrast, the black morph produced up to 14 capsules simultaneously, hatched an average of 178 veligers of 212 μm, and produced no nurse eggs. The intra-capsular development reached a more advanced stage in the brown-orange morph than in the black one. Several cytochrome subunit I (COI) and 16S rRNA (16S) haplotypes were found for the black morph, when compared to only one haplotype for both genes present for the brown-orange morph. Preliminary phylogenetic analyses separated the morphs into different clades, supported by robust bootstrap values and posterior probabilities (>98). Our results indicate that the morphs are two different species: the black morph was identified as Petaloconchus cf. varians and the brown-orange morph as a non-described species, Petaloconchus sp. (orange). The first is endemic to the Caribbean, while the second is potentially an introduced species.     

Ecological commonalities among pelagic fishes: comparison of freshwater ciscoes and marine herring and sprat

Systematic comparisons of the ecology between functionally similar fish species from freshwater and marine aquatic systems are surprisingly rare. Here, we discuss commonalities and differences in evolutionary history, population genetics, reproduction and life history, ecological interactions, behavioural ecology and physiological ecology of temperate and Arctic freshwater coregonids (vendace and ciscoes, Coregonus spp.) and marine clupeids (herring, Clupea harengus, and sprat, Sprattus sprattus). We further elucidate potential effects of climate warming on these groups of fish based on the ecological features of coregonids and clupeids documented in the previous parts of the review. These freshwater and marine fishes share a surprisingly high number of similarities. Both groups are relatively short-lived, pelagic planktivorous fishes. The genetic differentiation of local populations is weak and seems to be in part correlated to an astonishing variability of spawning times. The discrete thermal window of each species influences habitat use, diel vertical migrations and supposedly also life history variations. Complex life cycles and preference for cool or cold water make all species vulnerable to the effects of global warming. It is suggested that future research on the functional interdependence between spawning time, life history characteristics, thermal windows and genetic differentiation may profit from a systematic comparison of the patterns found in either coregonids or clupeids.     

Observations on shell form and its ecological significance in thaisid gastropods of the genus Lepsiella in New Zealand

This paper is concerned with the ecological significance of variation in shell form within the thaisid gastropod genus Lepsiella in New Zealand. Shell form has been investigated by measurement of shell height and breadth, aperture length and width, the diameters of consecutive whorls, apical angle, shell weight, and shell capacity, although in many cases shell height and shell breadth could not be measured because of erosion. L.albomarginata has been studied intensively at 4 stations in the South Island, and L. scobina less intensively at 6 stations in the North and South Islands. Comparisons of pairs of characters between stations have been tested by regression analysis and analysis of covariance where appropriate. Shells of L. albomarginata are relatively taller and narrower, and have a thicker wall, at a very sheltered station (Hakahaka Bay) that at more wave-exposed stations. L. scobina (sensu stricto), characterised by the presence of spiral ribs on the shell, exhibits less striking but comparable differences in shell shape. In laboratory tests in a tidal tank the thicker-shelled L. albomarginata from a sheltered station (Hakahaka Bay in Port Underwood) was much better able to resist attack by the shore crab Hemigrapsus edwardsi than was L. albomarginata from a nearby wave-exposed station (Whites Bay, near Cape Campbell, South Island). L. scobina from both stations was resistant to attack. H. edwardsi abounds at sheltered stations, but is missing from wave-exposed rock reefs such as those at Whites Bay, so that the ability to survive encounters with shore crabs is ecologically important to L. albomarginata inhabiting sheltered stations. L. scobina occupies a lower zone on the shore, where it is probably liable to encounter other more powerful predators. Its spiral ribs probably strengthen the shell. We do not know to what extent differences in shell form and thickness depend on environmental factors, and to what extent they originate genetically. Thin shells are associated with an abundance of mussels (Mytilus edulis ssp. aoteanus or Modiolus neozelanicus). There is an interesting possibility that a scarcity of mussels or other food caused by superior nonspecific predators might result in the production of better-protected Lepsiella.     

Effects of spermatophores on male and female monarch butterfly reproductive success

Summary I present the results of experiments designed to measure the effects of spermatophores produced by male monarch butterflies on male and female reproductive success. There was wide variation in the number of matings by captive males, suggesting the potential for strong sexual selection on males. Male lifespan was not affected by total number of matings, nor did it differ between males that were allowed to mate and those not exposed to females. Two effects of spermatophores on female behavior or fecundity are reported; (1) Females that received large spermatophores delayed remating longer than those receiving small ones. (2) Females allowed to mate several times laid more eggs than singly-mated females. The relative importance of these effects is discussed in relation to monarch mating patterns.     

Determining factors that influence the dispersal of a pelagic species: A comparison between artificial neural networks and evolutionary algorithms

Because of increasing transport and trade there is a growing threat of marine invasive species being introduced into regions where they do not presently occur. So that the impacts of such species can be mitigated, it is important to predict how individuals, particularly passive dispersers are transported and dispersed in the ocean as well as in coastal regions so that new incursions of potential invasive species are rapidly detected and origins identified. Such predictions also support strategic monitoring, containment and/or eradication programs. To determine factors influencing a passive disperser, around coastal New Zealand, data from the genus Physalia (Cnidaria: Siphonophora) were used. Oceanographic data on wave height and wind direction and records of occurrences of Physalia on swimming beaches throughout the summer season were used to create models using artificial neural networks (ANNs) and Na?ve Bayesian Classifier (NBC). First, however, redundant and irrelevant data were removed using feature selection of a subset of variables. Two methods for feature selection were compared, one based on the multilayer perceptron and another based on an evolutionary algorithm. The models indicated that New Zealand appears to have two independent systems driven by currents and oceanographic variables that are responsible for the redistribution of Physalia from north of New Zealand and from the Tasman Sea to their subsequent presence in coastal waters. One system is centred in the east coast of northern New Zealand and the other involves a dynamic system that encompasses four other regions on both coasts of the country. Interestingly, the models confirm, molecular data obtained from Physalia in a previous study that identified a similar distribution of systems around New Zealand coastal waters. Additionally, this study demonstrates that the modelling methods used could generate valid hypotheses from noisy and complicated data in a system about which there is little previous knowledge.     

Protein content of spermatophores in relation to monandry/polyandry in butterflies

The evolution of a mating system, and specifically mating frequency, is dependent on the costs and benefits to both sexes of mating once or several times. In butterflies, males transfer a spermatophore that contains both sperm and accessory gland products. Accessory gland substances contain nutrients which, in some species, females use to increase their reproductive output and longevity. Nutrients contained in these packaged ejaculates represent investment by males in reproduction. Consequently, the nutritional composition of spermatophores may vary depending on the mating system. There are two lines of arguments concerning the evolution of the nutrient content of ejaculates. One hypothesis argues that male nuptial gifts evolved in the context of certainty of paternity and ease of finding mates; thus spermatophores of polyandrous males (with lower certainty of paternity and greater ease of finding mates) should contain less protein than those of monandrous males, since more spermatophores are produced on average. The other hypothesis argues that polyandry evolved in the context of maximization of male transfer of nutrients to females, and hence spermatophores of polyandrous males should contain more protein than those of monandrous males. In an attempt to distinguish between these two hypotheses, we determined how protein content of ejaculates varied with the degree of polyandry in nine species of pierid and two species of satyrid butterflies. We found that both relative ejaculate mass and protein content increased with the degree of polyandry. Hence our results are consistent with the view that polyandry has evolved in the context of male transfer of nutrients to females, and provides another example of a male adaptation to multiple mating in butterflies.     

Reversal and transfer of spermatophores by Octopus vulgaris and O. hummelincki

Copulatory behavior in the octopus consists of a patterned series of movements whose individual functions are not well understood. Observations and experiments on mating in Octopus vulgaris and O. hummelincki from Bimini, Bahamas and Haiti were made over a number of years in the laboratory. The study reveals how the male octopus reverses its spermatophore prior to transferring it to the female. The male’s terminal organ (penis) extrudes the sperm end of the spermatophore as it exits from Needham’s sac up to the cap or filament thread and then, holding onto the cap thread, inserts the ejaculatory end of the spermatophore into the groove of the hectocotylus, thus reversing it. It is hypothesized that the primary function of the cap or filament thread, one part of the spermatophore, is to be a handle by which the reversal and transfer is accomplished. The siphon is merely a conduit through which the terminal organ functions and plays no role whatsoever in either the reversal of the spermatophore or in its insertion into the groove. The stimulus to which the male orients to find the groove of the hectocotylus is the apex of the retracted interbrachial membrane between the third and fourth right arms, which bears a fixed spatial relationship to the origin of the groove. A number of deviations from the successful transfer process, which represent loss of sperm from the reproductive process, are illustrated. Despite these deviations, the delicately balanced integration of the various movements serves to transfer the spermatophore to the female to ensure reproductive success.     

Role of sinking in diatom life-history cycles: ecological,evolutionary and geological significance

Rapid mass sinking of cells following diatom blooms, observed in lakes and the sea, is argued here to represent the transition from a growing to a resting stage in the life histories of these algae. Mass sinking is of survival value in those bloom diatoms that retain viability over long periods in cold, dark water but not in warm, nutrient-depleted surface water. Mechanisms for accelerating sinking speed of populations entering a resting or seeding mode are proposed. Previously unexplained features of diatom form and behaviour take on a new meaning in this context of diatom seeding strategies. Diatoms have physiological control over buoyancy as declining growth is accompanied by increasing sinking rates, where the frustule acts as ballast. Increased mucous secretion in conjunction with the cell protuberances characteristic of bloom diatoms leads to entanglement and aggregate formation during sinking; the sticky aggregates scavenge mineral and other particles during descent which further accelerates the sinking rate. Such diatom flocs will have sinking rates of 100 m d^-1 or more. This is corroborated by recent observations of mass phytoplankton sedimentation to the deep sea. This mechanism would explain the origin of marine snow flocs containing diatoms in high productivity areas and also the well-known presence of a viable deep sea flora. That mortality is high in such a seeding strategy is not surprising. A number of species-specific variables pertaining to size, morphology, physiology, spore formation and frustule dissolution rate will determine the sinking behaviour and thus control positioning of resting stages in the water column or on the bottom. It is argued that sinking behaviour patterns will be environmentally selected and that some baffling aspects of diatom form and distribution can be explained in this light. Rapid diatom sedimentation is currently believed to be mediated by zooplankton faecal pellets, particularly those of copepods. This view is not supported by recently published observations. I speculate that copepod grazing actually retards rather than accelerates vertical flux, because faecal pellets tend to be recycled within the surface layer by the common herbivorous copepods. Egestion of undigested food by copepods during blooms acts as a storage mechanism, as ungrazed cells are likely to initiate mass precipitation and depletion of the surface layer in essential elements. Unique features of diatoms are discussed in the light of their possible evolution from resting spores of other algae. An evolutionary ecology of pelagic bloom diatoms is deduced from behavioural and morphological characteristics of meroplanktonic and tychopelagic forms. Other shell-bearing protistan plankters share common features with diatoms. Similar life-history patterns are likely to be present in species from all these groups. The geological significance of mass diatom sinking in rapidly affecting transfer of biogenic and mineral particles to the sea floor is pointed out.     

Feeding selectivity and coexistence in two deposit-feeding gastropods

The diets of 2 ecologically similar deposit-feeding gastropods (Batillaria attramentaria and Cerithidea california) are described from 3 salt marsh localities in central California, USA. Gut contents revealed high similarity in dietary composition and a predominance of benthic diatoms. Sympatric and allopatric populations of the gastropods did not differ in diet. In both species, diatom food size increased with snail length with large B. attramentaria selecting longer diatom sizes than large C. californica. Size distributions of B. attramentaria differed in sympatric and allopatric populations. In sympatric populations, snail size classes which exhibited the greatest amounts of overlap in diatom size did not occur together. Effects of size-specific behavioral interactions in the gastropods may explain these differences.     

Shallow-water benthic and pelagic metabolism:

Pelagic primary production and benthic and pelagic aerobic metabolism were measured monthly at one site in the estuarine plume region of the nearshore continental shelf in the Georgia Bight. Benthic and water-column oxygen uptake were routinely measured and supplemented with seasonal measures of total carbon dioxide flux. Average respiratory quotients were 1.18:1 and 1.02:1 for the benthos and water column, respectively. Benthic oxygen uptake ranged from 1.23 to 3.41 g O₂ m^-2 d^-1 and totalled 756 g O₂ m^-2 over an annual period. Water column respiration accounted for 60% of total system metabolism. Turnover rates of organic carbon in sediment and the water column were 0.09 to 0.18 yr^-1 and 6.2 yr^-1, respectively. Resuspension appeared to control the relative amounts of organic carbon, as well as the sites and rates of organic matter degradation in the benthos and water column. Most of the seasonal variation in benthic and pelagic respiration could be explained primarily by temperature and secondarily by primary productivity. On an annual basis, the shelf ecosystem appeared to be heterotrophic; primary production was 73% of community metabolism, which was 749 g C m^-2 yr^-1. The timing of heterotrophic periods through the year appeared to be closely related to both river discharge and the periodicity of growth and death of marsh macrophytes in the adjacent estuary. The results of this study support the estuarine outwelling hypothesis of Odum (1968).This is Contribution No. 530 from the University of Georgia Marine Institute. This work was supported by the Georgia Sea Grant College Program maintained by the National Oceanic and Atmospheric Administration, US Department of Commerce     

Discreteness of pelagic faunal regions

A variety of statistical techniques were used to examine the discreteness of mesopelagic fish species assemblages in the western North Atlantic. Although faunal groups could be isolated, the results were more consistent with a gradation between faunas rather than abrupt change. Few species were regularly associated, thus suggesting that, while individual species may respond to environmental gradients, recurrent groups of species do not.     

Substratum relationships of some tropical Pacific intertidal gastropods

Among tropical Indo-West Pacific herbivorous intertidal gastropods, high intertidal neritids, and limpets and certain lower intertidal limpets are restricted to a narrow range of substratum types. Littorinids, planaxids, low intertidal neritids, most lower intertidal limpets, and the trochid Monodonta labio tolerate a wide variety of substrata. This pattern of substratum specialization is related to the overall adaptive strategies of the various groups: those with a large area of soft tissue in contact with the substratum (neritids and limpets) often specialize to a particular substratum, especially at high shore levels, while those in which the area of contact between animal and substratum is small do not. High intertidal neritids and limpets able to live on a wide range of substrata tend to be species of limited geographic distribution. It is concluded that, even within a tropically similar group of animals (raspers in this case) relevant dimensions of niche separation depend upon the structure of the environment as well as upon the structure of the animals themselves.     

Mitochondrial energetics of benthic and pelagic Antarctic teleosts

Antarctic fauna are highly adapted to the frigid waters of the Southern Ocean. This study describes the in vitro temperature sensitivity of oxygen consumption rates measured in liver mitochondria from the pelagic notothenioid Pleuragramma antarcticum between 5 and 35 °C. Oxygen fluxes were measured after the addition of millimolar levels of pyruvate, malate, succinate and glutamate (state II, LEAK) and saturating levels of ADP [state III, oxidative phosphorylation (OXPHOS)]. State III respiration significantly decreased above 18.7 °C. A comparison of the oxidative capacities among P. antarcticum and other notothenioids showed significant differences in state III respiration, where benthic species exhibited about 50 % lower rates than P. antarcticum. In addition, state III respiration rates normalized per milligram of mitochondrial protein of P. antarcticum were up to eight times higher than state III rates reported in the literature for other notothenioids. The comparatively high respiration rates measured in this study may be explained by our approach, which engaged both complexes I and II under conditions of oxidative phosphorylation. State III rates of independently activated complexes I and II were found to range from 42 to 100 % of rates obtained when both complexes were activated simultaneously in the same species. The remarkable tolerance of P. antarcticum OXPHOS toward warmer temperatures was unexpected for an Antarctic stenotherm and may indicate that thermal sensitivity of their mitochondria is not the driving force behind their stenothermy.     

Temperature relationships of some tropical Pacific intertidal gastropods

Tissue and environmental-temperature measurements were made using thermistors in Hawaii, Guam, Palau and Singapore in the summer of 1970. Limpets have a very limited ability to regulate body temperature, and several species possessed tissue temperatures greater than those of the substratum. Some neritids and the trochid Monodonta labio possessed tissue temperatures as much as 2.2° to 2.8°C below substratum temperature. Littorinids attached by a mucus film to the substratum were found to have relatively lower body temperatures than those attached by the foot. Among neritids there is a tendency for high intertidal species to be better temperature regulators than low intertidal species. A very important factor in the temperature regulation of intertidal snails appears to be the extent of the area of contact between the animal and the substratum, which is greatest in limpets and least (negligible) in littorinids. Evaporative cooling and light coloration of the shell play a role in certain species and groups.     

Relationships between intertidal zonation and circatidal rhythmicity in littoral gastropods

The presence and phase of circatidal rhythmicity was correlated with vertical zonation and other ecological factors. Ten species were studied in the field and in aktographs under controlled conditions in the laboratory. Retina plicata, Melanerita atramentosa, Bembicium nanum, Austrocochlea obtusa and Morula marginalba occupied mid- or upper-littoral zonations, and were subjected to regular tidal influence. They were active at high water and for a period after the ebb, possibly as these were the times of least desiccation. Each possessed a circatidal activity rhythm under non-tidal conditions. Midlittoral Amphinerita polita and lower-littoral Theliostyla albicilla were active at low water, possibly because their prefered habitats reduced desiccation and other selective forces, for example wave action and predation, determined the phase of the rhythmicity. Supra- and upper-littoral Nodilittorina pyramidalis and Melarapha unifasciata and lower- and infralittoral specimens of Bembicium auratum displayed no overt circatidal rhythmicity, possibly because they were not subjected to regular tidal action.     

Evolutionary ecology of settlement size in planktotrophic neritimorph gastropods

Body size during larval development is one of the most important attributes of aquatic animals. The optimal size for changing form or habitat may vary according to ecological traits of species, while phylogenetic constraints also play a significant role. The major goal of this study is to reveal the patterns in the settlement size of planktotrophic larvae in an archaic gastropod superorder Neritimorpha. We take advantage of the fact that size at various ontogenetic stages of neritimorphs can be rigorously estimated by measuring features of the adult opercula. This unique feature of neritimorphs has allowed us to generate the largest data set so far on larval settlement sizes within a group of marine invertebrates that recruit into very different post-metamorphic habitats. Eighty-eight species that represent most extant genera from rocky shores, seagrass beds, mangroves, estuaries, streams, submarine caves, deep-sea vents or seeps showed negligible intraspecific variation and considerable interspecific differences in settlement size, particularly between genera or families. Settlement size is determined primarily by phylogenetic constraints, while parallel evolution toward smaller sizes was shown to occur exclusively in four independent clades (two living and two extinct) of amphidromous snails with a marine larval period followed by a limnic adult phase. The smaller settlement size may possibly reduce the risk of being wafted away from the estuaries of their natal streams through less time achieving metamorphic competence, while ability to make occasional long-distance trips is retained by the presence of a sufficiently long delay period. This delay period also seems to obscure the possible correlation between settlement size and geographic distribution range of neritimorph species, both fully marine and amphidromous.     

Morphological patterns in high-intertidal gastropods: Adaptive strategies and their limitations

An analysis of shell form in several families of high-intertidal herbivorous gastropods has revealed the existence of morphological gradients. Among tropical littorinids, relative spire height and degree of development of external shell sculpture generally increase interspecifically, and often intraspecifically, from low to high shore levels. This gradient is also evident from temperate to tropical latitudes, and may be modified or overridden by exposure to wave action and other factors. Among acmaeid, patellid, and siphonariid limpets, there is a well-marked interspecific increase in relative shell height with increasing shore level, but latitudinal trends in morphology cannot at present be distinguished. Evidence from both tropical and temperate coasts indicates that limpets found in shaded places tend to have smoother shells than those often exposed to direct sunlight. In the Neritidae, relative shell globosity tends to increase interspecifically from low to high shore levels; most high-shore species have strong shell sculpture, with the base of the shell only slightly excavated. A decrease in the number of neritid species towards higher latitudes is accompanied by selective deletion of globose, strongly sculptured types living at high shore levels. Neritids and some limpets can employ evaporative cooling as a mechanism for temperature regulation. The morphological gradients in these groups reflect a relative increase in the volume of the extravisceral cavity (water reservoir) and a decrease in the area in contact with the substratum, as well as a relative decrease in the rate of water loss, in an upshore direction. Neritids are better adapted than limpets to high temperature and desiccation stresses because of their ability to regulate rate of evaporation and contact between soft parts and substratum. These differences are reflected in the complementary patterns of latitudinal diversity in the two groups. In Littorinidae, which attach to the substratum by a mucus film, the morphological gradients reflect a minimization of water loss and substratum contaet, and a maximization of reflective relative to absorptive surface area.This paper is an expanded version of part of a Ph. D. dissertation presented to Yale University.     

Heavy metal selection of phosphoglucose isomerase allozymes in marine gastropods

The effects of zinc and copper pollution on the allozymic variation of phosphoglucose isomerase (Pgi) genotypes were tested in the Mediterranean marine gastropods Monodonta turbinata and M. turbiformis in 26 laboratory tests involving 2 081 adult individuals, with 545 survivors. Our results indicate differential survivorship of electrophoretical Pgi allozyme genotypes for both pollutants. Zinc pollution selected against the allozyme genotype SS in M. turbinata, whereas no differential survivorship was observed in M. turbiformis which lacks this allozyme. Copper pollution selected against the MM genotype in both species. These results are inconsistent with the neutral theory of allozyme polymorphisms and appear to reflect the adaptive nature of some Pgi genotypes in these marine gastropods.