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ABSTRACT AimThuridillaBergh, 1872, is a lineage of herbivorous sea slugs externally distinguished by bright colours and distinctive patterns of lines and spots. Recent work revealed an exceptionally rapid, cryptic radiation of 13 species in the Indo‐Pacific, raising questions about mechanisms of speciation in this group. Here, we (i) study the diversification and historical biogeography ofThuridillain a phylogenetic context and (ii) assess the role of dispersal and vicariance as the predominant mode of speciation in the genus. LocationTropical and temperate regions of the Atlantic and Indo‐Pacific. Major Taxa StudiesGastropoda, Sacoglossa. MethodsA nearly complete taxon set with 28 out of 32 recognised species ofThuridillawas used, in a total sample of 172 specimens, together with sacoglossan outgroups. Phylogenetic relationships were determined using a multi‐locus approach combining two mitochondrial (COI and 16S) and one nuclear gene (H3). Species relationships, diversification times, and ancestral geographical ranges were inferred using relaxed‐clock methods together with Bayesian discrete phylogeographic methods under three calibration scenarios using the oldest known fossil of Sacoglossa,Berthelinia elegansCrosse, 1875, and tectonic events. ResultsThuridillaspecies branched off into four major clades in all calibration scenarios: two groups from the Atlantic plus Indo‐West Pacific (5 and 6 species) and two clades from the Indo‐West Pacific (4 and 17 species). The highest diversity of the genus is in the Western Pacific (14 spp.) with a peak in the East Indies Triangle (18 spp.), whereas the Atlantic is depauperate with only four species occurring in this ocean basin. Divergence between Atlantic and Indo‐West Pacific lineages occurred in two main temporal periods: the Miocene and the Pliocene. Speciation events within the 13 cryptic species‐complex fell mostly within Plio‐Pleistocene times. Main ConclusionsThe best supported hypothesis was an Indo‐West Pacific origin ofThuridillabetween 28 and 18 Mya during the Early Miocene. In the western Pacific, speciation likely occurred during transient allopatry during Plio‐Pleistocene sea‐level fluctuations. Under the three tested calibration scenarios, the limited diversity of the Atlantic Ocean is hypothesized to be derived from Miocene vicariant events associated with the closure of the Tethys Sea, dispersal across southern Africa, or long‐distance dispersal across the East Pacific Barrier prior to the uplift of the Isthmus of Panama.Thuridillais absent in the Eastern Pacific, potentially resulting from the extinction of ancestral lineages following the uplift of the Isthmus of Panama. Near‐complete sampling of diversity and reconstruction of historical biogeography thus yielded new insight into the relative contributions of dispersal versus vicariance during speciation over the history of this widely distributed, colourful genus. 

Abstract The discovery of bivalved sea slugs in the family Juliidae revolutionized our understanding of shell evolution in molluscs. However, relationships among lineages remain unclear, with generic names shared by extant and fossil taxa. Juliidae provide important fossil calibration points for dating phylogenies and for understanding the convergent evolution of a bivalved shell. Here, we present a revision of the systematics and classification of members of the Juliidae including Recent and fossil taxa by integrating molecular data from five genes, geometric morphometrics, and anatomical data. A list of all juliid taxa previously described is provided, along with diagnoses and remarks on genus-level names. Results from a geometric morphometric analysis support a new classification scheme, in which the genus Berthelinia, previously used for Recent and fossil taxa, is restricted to fossil species, and the genera Julia and Edenttellina are used exclusively for Recent species. Species of Edenttellina supported by molecular analyses are illustrated and anatomical details provided, including examination of protoconchs, radular teeth, and reproductive structures by scanning electron microscopy, along with illustrations of existing type material. Distinct biogeographical patterns in Julia and Edenttellina are discussed in the context of ecological differences between these two taxa, including host specificity and developmental mode. 

Abstract The genus Cyerce Bergh, 1870 has been a model for the study of defensive strategies, including chemical defences, ceratal autotomy, and crypsis or aposematism. Specialization on different algae and diverse genital armatures also make Cyerce a useful system for investigating speciation by host shift versus sexual selection. Here, we review the genus Cyerce in the Pacific and Indian oceans using molecular and morphological data. Two mitochondrial genes (COI and 16S) and one nuclear gene (H3) were sequenced from 154 specimens, including representatives from the Atlantic Ocean. Bayesian and maximum likelihood analyses were used to generate phylogenetic hypotheses. Species delimitation analyses performed on COI sequences recovered 17 genetically distinct Pacific and Indian Ocean species of Cyerce, 10 of which are new to science. Nine new species are named herein (C. takanoi sp. nov., C. katiae sp. nov., C. trowbridgeae sp. nov., C. blackburnae sp. nov., C. tutela sp. nov., C. basi sp. nov., C. whaapi sp. nov., C. goodheartae sp. nov., and C. liliuokalaniae sp. nov.). The 10th species, from the Red Sea, is not named due to the absence of internal anatomical data. These findings increase the species richness in Cyerce by about two-thirds, and demonstrate that even conspicuous taxa harbour considerable cryptic diversity. 

ABSTRACT Many coastal marine species experienced Pleistocene gene flow between the North Pacific and Atlantic. Understanding historical connectivity between ocean basins should aid in predicting how regional faunas will respond to recent warming that has intensified trans‐Arctic dispersal. Wetland fauna of the Northwestern Atlantic may have survived in estuarine refugia throughout glacial cycles, or recolonised from the southern coast, North Pacific or Northeastern Atlantic. Here, we used multilocus genetic markers and historical climate data to investigate lineage distribution and connectivity among populations of the nominally cosmopolitan sea slugAlderia modesta, sampled from mudflats on both coasts of the North Pacific and North Atlantic. Mitochondrial DNA clades from European and North American populations were deeply divergent and reciprocally monophyletic; differences at seven polymorphic nuclear loci indicated prolonged absence of trans‐Atlantic gene flow. A Pacific ancestor likely first colonised the Atlantic during the marine biotic interchange of the middle Pliocene ~3.5 Ma. Both mtDNA phylogenetics and nuclear genotype assignments support repeated trans‐Arctic colonisation of the Northwestern Atlantic from the Pacific during inter‐glacial cycles; no gene flow was evident since the last glacial maximum, however. Time‐calibrated coalescent phylogenies, Bayesian skyline plots and haplotype networks all indicated recent population expansions in the Pacific and Europe, but not Northwestern Atlantic. In both the Pacific and Northwestern Atlantic, older lineages persisted in patchy refugia north of glacial margins, while a derived clade of Pacific haplotypes indicates northward post‐LGM expansion. The biogeographical history ofAlderiacontrasts with rocky‐shore taxa that were largely extirpated by glacial advance and recolonised from refugia following the last glacial maximum. Based on molecular differences and distinctions in radular and penial stylet morphology, we resurrect the nameAlderia harvardiensisGould 1870 forAlderiafrom the Northwestern Atlantic and North Pacific;A. modestarefers exclusively to European slugs. 

Abstract Animals synthesize simple lipids using a distinct fatty acid synthase (FAS) related to the type I polyketide synthase (PKS) enzymes that produce complex specialized metabolites. The evolutionary origin of the animal FAS and its relationship to the diversity of PKSs remain unclear despite the critical role of lipid synthesis in cellular metabolism. Recently, an animal FAS-like PKS (AFPK) was identified in sacoglossan molluscs. Here, we explore the phylogenetic distribution of AFPKs and other PKS and FAS enzymes across the tree of life. We found AFPKs widely distributed in arthropods and molluscs (>6300 newly described AFPK sequences). The AFPKs form a clade with the animal FAS, providing an evolutionary link bridging the type I PKSs and the animal FAS. We found molluscan AFPK diversification correlated with shell loss, suggesting AFPKs provide a chemical defense. Arthropods have few or no PKSs, but our results indicate AFPKs contributed to their ecological and evolutionary success by facilitating branched hydrocarbon and pheromone biosynthesis. Although animal metabolism is well studied, surprising new metabolic enzyme classes such as AFPKs await discovery. 

Secondary metabolites often function as antipredator defenses, but when bioactive at low concentrations, their off-target effects on other organisms may be overlooked. Candidate “keystone molecules” are proposed to affect community structure and ecosystem functions, generally originating as defenses of primary producers; the broader effects of animal chemistry remain largely unexplored, however. Here, we characterize five previously unreported polyketides (alderenes A to E) biosynthesized by sea slugs reaching exceptional densities (up to 9000 slugs per square meter) in Northern Hemisphere estuaries. Alderenes comprise only 0.1% of slug wet weight, yet rendered live slugs or dead flesh unpalatable to three co-occurring consumers, making a potential food resource unavailable and redirecting energy flow in critical nursery habitat. Alderenes also displaced infauna from the upper sediment of the mudflat but attracted ovipositing snails. By altering communities, such compounds may have unexpected cascading effects on processes ranging from bioturbation to reproduction of species not obviously connected to the producing organisms, warranting greater attention by ecologists. 

Animal cytoplasmic fatty acid synthase (FAS) represents a unique family of enzymes that are classically thought to be most closely related to fungal polyketide synthase (PKS). Recently, a widespread family of animal lipid metabolic enzymes has been described that bridges the gap between these two ubiquitous and important enzyme classes: the animal FAS–like PKSs (AFPKs). Although very similar in sequence to FAS enzymes that produce saturated lipids widely found in animals, AFPKs instead produce structurally diverse compounds that resemble bioactive polyketides. Little is known about the factors that bridge lipid and polyketide synthesis in the animals. Here, we describe the function of EcPKS2 fromElysia chlorotica, which synthesizes a complex polypropionate natural product found in this mollusc. EcPKS2 starter unit promiscuity potentially explains the high diversity of polyketides found in and among molluscan species. Biochemical comparison of EcPKS2 with the previously described EcPKS1 reveals molecular principles governing substrate selectivity that should apply to related enzymes encoded within the genomes of photosynthetic gastropods. Hybridization experiments combining EcPKS1 and EcPKS2 demonstrate the interactions between the ketoreductase and ketosynthase domains in governing the product outcomes. Overall, these findings enable an understanding of the molecular principles of structural diversity underlying the many molluscan polyketides likely produced by the diverse AFPK enzyme family.