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The causes and consequences of genome reduction in animals are unclear because our understanding of this process mostly relies on lineages with often exceptionally high rates of evolution. Here, we decode the compact 73.8-megabase genome of Dimorphilus gyrociliatus, a meiobenthic segmented worm. The D. gyrociliatus genome retains traits classically associated with larger and slower-evolving genomes, such as an ordered, intact Hox cluster, a generally conserved developmental toolkit and traces of ancestral bilaterian linkage. Unlike some other animals with small genomes, the analysis of the D. gyrociliatus epigenome revealed canonical features of genome regulation, excluding the presence of operons and trans-splicing. Instead, the gene-dense D. gyrociliatus genome presents a divergent Myc pathway, a key physiological regulator of growth, proliferation and genome stability in animals. Altogether, our results uncover a conservative route to genome compaction in annelids, reminiscent of that observed in the vertebrate Takifugu rubripes.

Members of the following marine annelid families are found almost exclusively in the interstitial environment and are highly adapted to move between sand grains, relying mostly on ciliary locomotion: Apharyngtidae n. fam., Dinophilidae, Diurodrilidae, Nerillidae, Lobatocerebridae, Parergodrilidae, Polygordiidae, Protodrilidae, Protodriloididae, Psammodrilidae and Saccocirridae. This article provides a review of the evolution, systematics, and diversity of these families, with the exception of Parergodrilidae, which was detailed in the review of Orbiniida by Meca, Zhadan, and Struck within this Special Issue. While several of the discussed families have previously only been known by a few described species, recent surveys inclusive of molecular approaches have increased the number of species, showing that all of the aforementioned families exhibit a high degree of cryptic diversity shadowed by a limited number of recognizable morphological traits. This is a challenge for studies of the evolution, taxonomy, and diversity of interstitial families as well as for their identification and incorporation into ecological surveys. By compiling a comprehensive and updated review on these interstitial families, we hope to promote new studies on their intriguing evolutionary histories, adapted life forms and high and hidden diversity.

BACKGROUND: Diuronotus is one of the most recently described genera of Paucitubulatina, one of the three major clades in Gastrotricha. Its morphology suggests that Diuronotus is an early branch of Paucitubulatina, making it a key taxon for understanding the evolution of this morphologically understudied group. Here we test its phylogenetic position employing molecular data, and provide detailed descriptions of the muscular, nervous, and ciliary systems of Diuronotus aspetos, using immunohistochemistry and confocal laser scanning microscopy.RESULTS: We confirm the proposed position of D. aspetos within Muselliferidae, and find this family to be the sister group to Xenotrichulidae. The muscular system, revealed by F-actin staining, shows a simple, but unique organization of the trunk musculature with a reduction to three pairs of longitudinal muscles and addition of up to five pairs of dorso-ventral muscles, versus the six longitudinal and two dorso-ventral pairs found in most Paucitubulatina. Using acetylated α-tubulin immunoreactivity, we describe the pharynx in detail, including new nervous structures, two pairs of sensory cilia, and a unique canal system. The central nervous system, as revealed by immunohistochemistry, shows the general pattern of Gastrotricha having a bilobed brain and a pair of ventro-longitudinal nerve cords. However, in addition are found an anterior nerve ring, several anterior longitudinal nerves, and four ventral commissures (pharyngeal, trunk, pre-anal, and terminal). Two pairs of protonephridia are documented, while other Paucitubulatina have one. Moreover, the precise arrangement of multiciliated cells is unraveled, yielding a pattern of possibly systematic importance.CONCLUSION: Several neural structures of Diuronotus resemble those found in Xenotrichula (Xenotrichulidae) and may constitute new apomorphies of Paucitubulatina, or even Gastrotricha. In order to test these new evolutionary hypotheses, comparable morphologi

Spiralia is a vast clade of Metazoa comprising large and well-known organisms, e.g., Annelida and Mollusca, but also many microscopic animals such as Gastrotricha or Gnathifera (including, Rotifera) of the often overlooked meiofauna. To date, the phylogeny and morphology of Spiralia have been difficult to resolve and understand. The present thesis focuses on spiralian meiofauna to i) reconstruct the phylogeny of this clade using transcriptomics and place enigmatic meiofaunal taxa and ii) resolve the morphology of three important taxa, mainly employing confocal laser scanning microscopy and immunohistochemistry: the spiralian incertae sedis, Lobatocerebrum, the recently described monospecific phylum Micrognathozoa (Gnathifera), and an early branching Gastrotricha, Diuronotus aspetos. The new spiralian phylogeny reveals with high support that the deepest branches of Spiralia consist of meiofaunal representatives, that Gnathifera is the sister group of remaining Spiralia, that Gastrotricha+Platyhelminthes branches off next and that Lobatocerebrum is an Annelida. The morphological surveys of the musculature, nervous system, glands, and ciliation on three phylogenetically distinct taxa yield more insight into their evolution: Lobatocerebrum is an aberrant annelid showing only few common traits with Annelida, yet, our detailed studies unravel putative resemblances of muscular, nervous and glandular system to previous findings in annelids. Micrognathozoa shows more resemblances with Rotifera than Gnathostomulida (these three taxa together forming Gnathifera). Furthermore, we could infer possible plesiomorphic states of Gnathifera such as the paired ventro-lateral nerve chords (shared with many Spiralia) as well as recover putative Gnathifera apomorphies such as the pharyngeal ganglion; all adding new information on the evolution of this group. Diuronotus aspetos shows a unique combination o