Onuphid polychaetes are tubicolous marine worms commonly reported worldwide from intertidal areas to hadal depths. They often dominate in benthic communities and have economic importance in aquaculture and recreational fishing. Here we report the phylogeny of the family Onuphidae based on the combined analyses of nuclear (18S rDNA) and mitochondrial (16S rDNA) genes. Results of Bayesian and Maximum Likelihood analyses supported the monophyly of Onuphidae and its traditional subdivision into two monophyletic subfamilies: Onuphinae and Hyalinoeciinae. Ten of 22 recognized genera were monophyletic with strong node support; four more genera included in this study were either monotypic or represented by a single species. None of the genera appeared para- or polyphyletic and this indicates a strong congruence between the traditional morphology-based systematics of the family and the newly obtained molecular-based phylogenetic reconstructions. Intergeneric relationships within Hyalinoeciinae were not resolved. Two strongly supported monophyletic groups of genera were recovered within Onuphinae: ((Onuphis, Aponuphis), Diopatra, Paradiopatra) and (Hirsutonuphis, (Paxtonia, (Kinbergonuphis, Mooreonuphis))). A previously accepted hypothesis on the subdivision of Onuphinae into the Onuphis group of genera and the Diopatra group of genera was largely rejected.
In this study, we analyze the current state of knowledge on extant Eunicida systematics, morphology, feeding, life history, habitat, ecology, distribution patterns, local diversity and exploitation. Eunicida is an order of Errantia annelids characterized by the presence of ventral mandibles and dorsal maxillae in a ventral muscularized pharynx. The origin of Eunicida dates back to the late Cambrian, and the peaks of jaw morphology diversity and number of families are in the Ordovician. Species richness is heterogeneous among the seven recent families, with more than half of the valid species belonging to the Eunicidae + Onuphidae clade, one of the latest clades to diverge. Eunicidans inhabit soft and hard substrates from intertidal to deep waters in all oceans. The few freshwater species are restricted to Histriobdellidae, a family exclusively commensal/parasite of crustaceans. The reproductive biology, development and ecology of most families are poorly known and the information available suggests low dispersal ability. However, all families have records of widely distributed species. Scrutiny of these wide distributions has often revealed the presence of exotic species or more than one species. The exploration of the deep-sea and of new habitats has led to recent descriptions of new species. Furthermore, the revision of type specimens, the examination of new morphological features and the use of molecular data have revealed hidden biodiversity under unjustified synonyms, poor understanding of morphological features and incomplete descriptions. Molecular studies are still very few or nonexistent for the families Histriobdellidae, Hartmaniellidae, Lumbrineridae and Oenonidae. The integration of new methodologies for morphological and molecular study, along with information on biological and ecological traits appears to be the path to improve the knowledge on the diversity of Eunicida.
Background: Modern integrative taxonomy-based annelid species descriptions are detailed combining morphological data and, since the last decades, also molecular information. Historic species descriptions are often comparatively brief lacking such detail. Adoptions of species names from western literature in the past led to the assumption of cosmopolitan ranges for many species, which, in many cases, were later found to include cryptic or pseudocryptic lineages with subtle morphological differences. Natural history collections and databases can aid in assessing the geographic ranges of species but depend on correct species identification. Obtaining DNA sequence information from wet-collection museum specimens of marine annelids is often impeded by the use of formaldehyde and/or long-term storage in ethanol resulting in DNA degradation and cross-linking. Results: The application of ancient DNA extraction methodology in combination with single-stranded DNA library preparation and target gene capture resulted in successful sequencing of a 110-year-old collection specimen of quill worms. Furthermore, a 40-year-old specimen of quill worms was successfully sequenced using a standard extraction protocol for modern samples, PCR and Sanger sequencing. Our study presents the first molecular analysis of Hyalinoecia species including the previously known species Hyalinoecia robusta, H. tubicloa, H. artifex, and H. longibranchiata, and a potentially undescribed species from equatorial western Africa morphologically indistinguishable from H. tubicola. The study also investigates the distribution of these five Hyalinoecia species. Reassessing the distribution of H. robusta reveals a geographical range covering both the Atlantic and the Indian Oceans as indicated by molecular data obtained from recent and historical specimens. Conclusion: Our results represent an example of a very wide geographical distribution of a brooding deep-sea annelid with a complex reproduction strategy and seemingly very limited dispersal capacity of its offspring, and highlights the importance of molecular information from museum specimens for integrative annelid taxonomy and biogeography.
Diopatra Audouin & Milne-Edwards, 1833 is a species rich genus that is common in tropical and subtropical regions. The genus is readily identified by its striking, spiral branchiae, but species identification has historically been challenging due to a high variation in diagnostic characters used. This study aims to reconstruct the phylogeny of Diopatra with molecular markers and assess the species diversity of West African Diopatra with the species delimitation programs bPTP and BPP. Specimens were collected from Morocco to Angola, and the markers COI, 16S and 28S were sequenced from 76 specimens. The constructed phylogeny retrieved Diopatra as monophyletic, as well as five well supported clades within the genus. All clades were defined by morphological characters, some of which have previously not been considered to have high phylogenetic or taxonomical value. Species delimitation analyses recovered 17 new species, several of which were not readily identified morphologically. One species complex comprising between one and 12 species was left unresolved due to incongruence between the species delimitation methods and challenging morphology. Our results indicate that the diversity of Diopatra is significantly underestimated, where this regional study near to doubled the number ofknown species from the East Atlantic.
Lumbrineridae is a family of marine annelids with simple external morphology but complex and diverse jaw apparatuses consisting of paired maxillae and mandibles. Here we present the first phylogeny of lumbrinerids based on combination of nuclear (18S rDNA) and mitochondrial (COI, 16S rDNA) markers utilizing Bayesian inference and Maximum Likelihood approaches. Despite limited taxon sampling, our results support monophyly of the genera Abyssoninoe, Augeneria, Gallardoneris, Lumbrineriopsis, and Ninoe and indicate polyphyly of the genera Lumbrineris (the type genus of the family) and Scoletoma. None of the morphological characters traditionally used in lumbrinerid systematics, such as the presence of connecting plates, four pairs of maxillae, bidentate simple hooded hooks, colorless maxillae IV, and multidentate maxillae IV were found to be exclusive homologies for a well-supported clade and have probably evolved several times independently within Lumbrineridae.