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AbstractMost organisms are faced with daily cyclic changes in a suite of environmental conditions, including temperature. In shallow marine waters, populations of the same species may experience either intertidal or subtidal conditions, such that some individuals experience extreme daily fluctuations as the tide ebbs and flows, while others only a few meters away experience less pronounced or less frequent fluctuations or almost constant thermal conditions. This study used a fully factorial combination of three thermal treatments and two diet treatments to test the hypotheses that (1) individuals experiencing fluctuating temperatures perform more poorly than those experiencing the same mean temperature under constant conditions and that (2) the negative impact of fluctuating temperatures is greater under food-limiting conditions. Five life-history components of the slipper snail Crepidula cf. marginalis were used as response variables. We found that temperature fluctuations impacted size at hatching and time to hatching, as well as growth rate, to some extent. Diet treatments impacted growth rates, clutch sizes, time to first brood, and time to hatching. There were no statistically significant interactions between the two factors. These results show that fluctuations between two temperatures that are typically experienced by these animals in the field can significantly affect fitness-related characters and, therefore, suggest the tidal height at which larvae settle can significantly impact individual fitness. This is one of the first studies to demonstrate that differences in the frequency of fluctuations, in the absence of differences in the magnitude or the mean, can have significant impacts on invertebrate life histories.

Biodiversity assessments are critical for setting conservation priorities, understanding ecosystem function and establishing a baseline to monitor change. Surveys of marine biodiversity that rely almost entirely on sampling adult organisms underestimate diversity because they tend to be limited to habitat types and individuals that can be easily surveyed. Many marine animals have planktonic larvae that can be sampled from the water column at shallow depths. This life stage often is overlooked in surveys but can be used to relatively rapidly document diversity, especially for the many species that are rare or live cryptically as adults. Using DNA barcode data from samples of nemertean worms collected in three biogeographical regions-Northeastern Pacific, the Caribbean Sea and Eastern Tropical Pacific-we found that most species were collected as either benthic adults or planktonic larvae but seldom in both stages. Randomization tests show that this deficit of operational taxonomic units collected as both adults and larvae is extremely unlikely if larvae and adults were drawn from the same pool of species. This effect persists even in well-studied faunas. These results suggest that sampling planktonic larvae offers access to a different subset of species and thus significantly increases estimates of biodiversity compared to sampling adults alone. Spanish abstract is available in the electronic supplementary material.

AbstractThe identity of wild cloning sea star larvae has been a mystery since they were first documented in the Caribbean. The most commonly collected cloning species was thought to belong to the Oreasteridae, on the basis of similarity with sequences from Oreaster reticulatus and Oreaster clavatus . This larval form has recently been linked to a rare benthic juvenile. As part of two larger DNA barcoding projects, we collected cloning asteroid larvae from the Caribbean coast of Panama and compared them to a large reference database of tropical echinoderms. Morphological and DNA barcode data from the cytochrome c oxidase subunit I gene demonstrated that Panamanian larvae belonged to the same operational taxonomic unit as those recovered in previous studies of cloning larvae from the Caribbean. Much to our surprise, sequences from these larvae clearly identified them as belonging to Valvaster striatus , a species typically considered to be endemic to the Indo-West Pacific. A lineage of Mithrodia clavigera that occurs in both the Caribbean and the Indo-West Pacific also has cloning larvae, suggesting that this unusual life history has allowed larvae to pass around the Cape of Good Hope and the Benguela upwelling region, which is a barrier to dispersal for most tropical marine invertebrates.

In species with complex life cycles, early developmental stages are often less thermally tolerant than adults, suggesting that they are key to predicting organismal response to environmental warming. Here we document the optimal and lethal temperatures of larval sea urchins, and we use those to calculate the warming tolerance and the thermal safety margin of early larval stages of seven tropical species. Larvae of Echinometra viridis, Echinometra lucunter, Lytechinus williamsi, Eucidaris tribuloides, Tripneustes ventricosus, Clypeaster rosaceus, and Clypeaster subdepressus were reared at 26, 28, 30, 32, and 34 °C for 6 days. The temperatures at which statistically significant reductions in larval performance are evident are generally the same temperatures at which statistically significant reductions in larval survival were detected, showing that the optimal temperature is very close to the lethal temperature. The two Echinometra species had significantly higher thermal tolerance than the other species, with some surviving culture temperatures of 34 °C and showing minimal impacts on growth and survival at 32 °C. In the other species, larval growth and survival were depressed at and above 30 or 32 °C. Overall, these larvae have lower warming tolerances (1 to 5 °C) and smaller thermal safety margins (-3 to 3 °C) than adults. Survival differences among treatments were evident by the first sampling on day 2, and survival at the highest temperatures increased when embryos were exposed to warming after spending the first 24 hours at ambient temperature. This suggests that the first days of development are more sensitive to thermal stress than are later larval stages.

Chemical signaling, especially signaling with waterborne cues, is an important mode of communication between conspecifics of aquatic organisms. Although conspecific associations play an important role in sex allocation of sequential hermaphroditic slipper limpets, the mode of signaling is unknown. We tested the hypothesis that the effects of conspecifics on animal size and time of sex change in the tropical slipper limpet Crepidula cf. marginalis are mediated by waterborne cues. In our experiment, pairs of snails (one small and one large) were kept in cups, either together or partitioned off with fine or coarse mesh, or partitioned, but switched from side to side to allow contact with the cup mate's pedal mucus. The larger snails that were allowed contact with the smaller companions grew faster, and generally changed sex sooner, than did the larger snails in the barrier treatments, which allowed no physical contact. The smaller snails that were allowed contact with the larger cup mate delayed sex change compared to those separated from their cup mates. We were, therefore, able to reject the hypothesis that waterborne cues mediate communication between these snails. Our results suggest that the cue that affects size and time to sex change requires some kind of physical interaction that is lost when the snails are separated. Furthermore, contact with another snail's pedal mucus does not compensate for the loss of physical contact. Since males often attach to the shell of larger females, direct contact may mediate this kind of physical interaction via positional information, physical stimulation, or contact-based chemical communication. Whatever the cue, contact with conspecifics influences both partners, resulting in, surprisingly, a higher growth rate in the larger animal and delayed sex change in the smaller animal., (© 2015 Marine Biological Laboratory.)

Ocean Sampling Day was initiated by the EU-funded Micro B3 (Marine Microbial Biodiversity, Bioinformatics, Biotechnology) project to obtain a snapshot of the marine microbial biodiversity and function of the world's oceans. It is a simultaneous global mega-sequencing campaign aiming to generate the largest standardized microbial data set in a single day. This will be achievable only through the coordinated efforts of an Ocean Sampling Day Consortium, supportive partnerships and networks between sites. This commentary outlines the establishment, function and aims of the Consortium and describes our vision for a sustainable study of marine microbial communities and their embedded functional traits.

Molluscs show a wide diversity of sexual systems and strategies. There are both gastropod and bivalve families that are each primarily dioecious, simultaneous hermaphrodites, or sequential hermaphrodites, and other families in which almost every sexual strategy occurs. The multiple evolutionary transitions of sexual systems within molluscs would allow comparative analyses of the associated ecological factors, but data on all but a few groups are too sparse to draw many solid conclusions. The phylogenetic distribution of sexual systems in the Mollusca shows that gastropods and bivalves demonstrate different patterns, possibly associated with the presence/absence of copulation. The distribution of change of sex suggests that, in gastropods, sequential hermaphrodites do not evolve from simultaneous hermaphrodites, and that sex reversal (flip-flopping) occurs in free-spawners but not in copulators. Three well-studied protandrous gastropod groups (calyptraeids, coralliophilids, and patellogastropods) show similar responses to environmental conditions and associations with conspecifics. They all have the following attributes: (1) they are sedentary, (2) they live in groups, patches, or aggregates, and (3) size at sex change varies among sites and among aggregates. In addition the available experimental evidence suggests that (4) the presence of females or large individuals represses growth and sex change of males, and (5) behavior seems to mediate the repressive influence of large females. Available data from other species tend to support these patterns. Finally, the repression of growth of males by females in protandry likely facilitates the evolution of dwarf males.

Mode of development in marine invertebrates has been largely viewed as a dichotomy between small eggs that develop into free-living planktotrophic larvae and large eggs that bypass the larval stage and develop directly into juveniles. Modes of development that could be categorized as "intermediate" between these two extremes include facultative feeding larvae, short-lived planktotrophic larvae, dispersal dimorphisms, and poecilogony (in which nutritional mode varies within a species). These intermediates are rare. The few species-level phylogenies available that include them do not generally support the interpretation of intermediates as necessary, ephemeral transitional forms. Instead, they support the idea that intermediates are well adapted to their environments but that either these environments are short-lived relative to the frequency of speciation, or speciation events are associated with shifts in the mode of development. Each of the different intermediate forms could have evolved in response to variable environments. The phenotypically plastic intermediates could be a response to predictable environmental variation. Facultative feeding larvae and short-lived planktotrophic larvae could reflect conservative bet-hedging in response to unpredictable environmental variation, whereas poecilogony with mixed clutches could represent the alternative: diversifying bet-hedging. Since environmental variability is common, it remains an enigma why these intermediates are so rare. Discovery of more intermediates, and their careful description in terms of the level of variation expressed within, and among, clutches, and among females and populations, as well as determination of the genetic and environmental influences on this variation, will provide valuable test-cases for theories of the evolution of alternative phenotypes.

Gastropods have lost the quintessential snail feature, the coiled shell, numerous times in evolution. In many cases these animals have developed a limpet morphology with a cap-shaped shell and a large foot. Limpets thrive in marginal habitats such as hydrothermal vents, the high-energy rocky intertidal areas and fresh water, but they are considered to be evolutionary dead-ends, unable to re-evolve a coiled shell and therefore unable to give rise to the diversity seen among coiled snails. The re-evolution of a coiled shell, or any complex character, is considered unlikely or impossible (Dollo's law) because the loss of the character is followed by the loss of the genetic architecture and developmental mechanisms that underlie that character. Here, we quantify the level of coiling in calyptraeids, a family of mostly uncoiled limpets, and show that coiled shells have re-evolved at least once within this family. These results are the first demonstration, to our knowledge, of the re-evolution of coiling in a gastropod, and show that the developmental features underlying coiling have not been lost during 20-100 Myr of uncoiled evolutionary history. This is the first example of the re-evolution of a complex character via a change in developmental timing (heterochrony) rather than a change in location of gene expression (heterotopy).