Understanding the ecological and evolutionary forces that determine the genetic structure and spread of invasive species is a key component of invasion biology. The bay barnacle, Balanus improvisus (= Amphibalanus improvisus), is one of the most successful aquatic invaders worldwide, and is characterised by broad environmental tolerance. Although the species can spread through natural larval dispersal, human-mediated transport through (primarily) shipping has almost certainly contributed to the current global distribution of this species. Despite its worldwide distribution, little is known about the phylogeography of this species. Here, we characterize the population genetic structure and model dispersal dynamics of the barnacle B. improvisus, and describe how human-mediated spreading via shipping as well as natural larval dispersal may have contributed to observed genetic variation. We used both mitochondrial DNA (cytochrome c oxidase subunit I: COI) and nuclear microsatellites to characterize the genetic structure in 14 populations of B. improvisus on a global and regional scale (Baltic Sea). Genetic diversity was high in most populations, and many haplotypes were shared among populations on a global scale, indicating that long-distance dispersal (presumably through shipping and other anthropogenic activities) has played an important role in shaping the population genetic structure of this cosmopolitan species. We could not clearly confirm prior claims that B. improvisus originates from the western margins of the Atlantic coasts; although there were indications that Argentina could be part of a native region. In addition to dispersal via shipping, we show that natural larval dispersal may play an important role for further colonisation following initial introduction.

Pygospio elegans is an opportunistic, wide-spread spionid polychaete that reproduces asexually via fragmentation and can produce benthic and pelagic larvae, hence combining different developmental modes in one species. We documented the density, size distribution, and reproductive activity of P. elegans at four sites in the Danish Isefjord-Roskilde Fjord estuary complex, where all modes of reproduction were reported. We compared population dynamics of this species to environmental parameters such as salinity, temperature, and sediment characteristics (grain size, sorting, porosity, water content, organic content, C/N). We observed that new cohorts-resulting either from sexual or asexual reproduction-appeared in spring and fall, and old ones disappeared in late summer and winter. Sexual reproduction occurred from September until May, and although their timing was variable, there were two reproductive peaks at three sites. At those sites, we also observed a switch in larval developmental mode. Asexual reproduction peaked in April. While the seasonal dynamics can be related to temperature to a large extent, the differences in population dynamics among sites also correlated with sediment structure and salinity. Populations from sites with coarse and heterogeneous sediment had high levels of sexual reproduction. At the site with lower salinity, intermediate and benthic larvae were present during winter in contrast to pelagic larvae found at the other sites. However, we could not identify one clear environmental factor determining the mode of development. At present, it remains unclear to what degree genetic background contributes to mode of development. Hence, whether the differences in developmental mode are the result of genetically different cohorts will be further investigated.Additional key words: life history, poecilogony, development, spatiotemporal variation environmental impact Invertebrate Biology 135(4): 370-384.

BackgroundHaemocyanin is the respiratory protein of most of the Mollusca. In cephalopods and gastropods at least two distinct isoforms are differentially expressed. However, their physiological purpose is unknown. For the common cuttlefish Sepia officinalis, three isoforms are known so far, whereas for only two of them the complete mRNA sequences are available. In this study, we sequenced the complete mRNA of the third haemocyanin isoform and measured the relative expression of all three isoforms during embryogenesis to reveal a potential ontogenetic relevance.ResultsThe cDNA of isoform 3 clearly correlates to the known Sepia officinalis haemocyanin subunits consisting of eight functional units and an internal duplicated functional unit d. Our molecular phylogenetic analyses reveal the third isoform representing a potentially ancestral haemocyanin isoform, and the analyses of the expression of haemocyanin type 3 reveal that haemocyanin type 3 only can be observed within eggs and during early development. Isoforms 1 and 2 are absent at these stages. After hatching, isoform 3 is downregulated, and isoform 1 and 2 are upregulated.ConclusionsOur study clearly shows an embryonic relevance of the third isoform, which will be further discussed in the light of the changes in the physiological function of haemocyanin during ontogeny. Taken together with the fact that it could also be the isoform closest related to the common ancestor of cuttlefish haemocyanin, the phylogeny of cuttlefish haemocyanin may be recapitulated during its ontogeny.

Interactions between symbiotic organisms and their hosts dramatically influence not only organismal ecology and evolution, but also the dynamics of entire ecosystems (Faust & Raes, 2012;Godfrey-Smith, 2015).For the majority of symbioses, however, we do not know how the interacting species affect each other, much less the broader consequences of the symbiosis for communities or ecosystems. Often, it is the lack of tools for effectively monitoring or investigating symbiotic interactions that prohibits progress. In addition, a lack of general knowledge