The phylogenetic relationships of selected members of the phylum Nemertea are explored by means of six markers amplified from the genomic DNA of freshly collected specimens (the nuclear 18S rRNA and 28S rRNA genes, histones H3 and H4, and the mitochondrial genes 16S rRNA and cytochrome c oxidase subunit I). These include all previous markers and regions used in earlier phylogenetic analyses of nemerteans, therefore acting as a scaffold to which one could pinpoint any previously published study. Our results, based on analyses of static and dynamic homology concepts under probabilistic and parsimony frameworks, agree in the non‐monophyly of Palaeonemertea and in the monophyly of Heteronemerta and Hoplonemertea. The position of Hubrechtella and the Pilidiophora hypothesis are, however, sensitive to analytical method, as is the monophyly of the non‐hubrechtiid palaeonemerteans. Our results are, however, consistent with the main division of Hoplonemertea into Polystilifera and Monostilifera, the last named being divided into Cratenemertea and Distromatonemertea, as well as into the main division of Heteronemertea into Baseodiscus and the remaining species. The study also continues to highlight the deficient taxonomy at the family and generic level within Nemertea and sheds light on the areas of the tree that require further refinement. © The Willi Hennig Society 2011.
Resolving the deep relationships of ancient animal lineages has proven difficult using standard Sanger-sequencing approaches with a handful of markers. We thus reassess the relatively well-studied phylogeny of the phylum Nemertea (ribbon worms)-for which the targeted gene approaches had resolved many clades but had left key phylogenetic gaps-by using a phylogenomic approach using Illumina-based de novo assembled transcriptomes and automatic orthology prediction methods. The analysis of a concatenated data set of 2,779 genes (411,138 amino acids) with about 78% gene occupancy and a reduced version with 95% gene occupancy, under evolutionary models accounting or not for site-specific amino acid replacement patterns results in a well-supported phylogeny that recovers all major accepted nemertean clades with the monophyly of Heteronemertea, Hoplonemertea, Monostilifera, being well supported. Significantly, all the ambiguous patterns inferred from Sanger-based approaches were resolved, namely the monophyly of Palaeonemertea and Pilidiophora. By testing for possible conflict in the analyzed supermatrix, we observed that concatenation was the best solution, and the results of the analyses should settle prior debates on nemertean phylogeny. The study highlights the importance, feasibility, and completeness of Illumina-based phylogenomic data matrices.
The life history of unionid bivalve molluscs includes retention of developing embryos within the gills of parental mussels. This brooding behavior may facilitate nutrient transfer to the glochidia larvae, i.e., matrotrophy. To address this possibility, morphological relationships between brood chambers and developing larvae of Pyganodon cutaructu and Utterbackia imbecillis were examined with TEM, and larval shells were observed with SEM, for features that could be associated with the uptake of dissolved materials. Early in brooding, glochidia are enclosed in a vitelline membrane that physically contacts numerous cilia and microvilli of the epithelial cells lining the brood chamber (marsupium). The vitelline membrane subsequently disappears. Lamellar tissues of parental mussels initially have large deposits of glycogen that diminish during the course of brooding. Septa separating brood chambers from adjacent secondary water tubes have numerous mitochondria and microvilli, suggesting the potential for active transport of materials into or out of the marsupia. Since punctae (pores) in the larval shells become filled with an organic matrix early in brooding, they are unlikely to be involved in nutrient exchange. Ultrastructnre of the brood chamber and physical contact between the parental mussel and larvae are consistent with a nutritive role for retention of glochidia in the marsupia.
The heteronemertean genus Dushia Corrêa, 1963 was established for what was identified as D. atra (Girard, 1851) (originally Meckelia atra) based on material from littoral, shallow waters in Curaçao, while the nominal species Meckelia atra was originally described from deep water off Florida Cape. In this paper, we conclude that the type species for Dushia has been misidentified. Based on specimens from the Caribbean, we establish D. wijnhoffae Schwartz & Norenburg sp. nov. to represent the true identity of the genus, according to Article 70.3.2 of the International Code of Zoological Nomenclature; Meckelia atra should be regarded as a nomen dubium. While the genus has remained monotypic since its establishment, our molecular analysis discovered a second member—or rather a group of members—from the West Pacific. This ‘group of members’, herein termed Dushia nigra (Stimpson, 1855) species complex comb. nov., involves i) at least two genetically separated biological entities, 0.136–0.148 (p-distance) and 0.152–0.168 (K2P) apart in terms of 513-bp COI sequences, which we interpret as likely to represent cryptic species, ii) three color forms, orange, brown, and black, with the last one occurring most frequently, and iii) four nominal species, Meckelia nigra Stimpson, 1855 (now Cerebratulus niger), Meckelia rubella Stimpson, 1855 (now Cerebratulus rubellus), Micrura formosana Yamaoka, 1939, and Micrura japonica Iwata, 1952. At present, however, we have no objective ground as to which of the four potentially available names (i.e., formosana, japonica, nigra, and rubella) should be allotted to the two cryptic species discovered in the analysis, because i) a single locality can harbor two cryptic species, ii) a single cryptic species may contain three different color morphs (i.e., orange, brown, black), and iii) no data from the type localities for these four nominal species are available at the moment. Our multi-locus analysis of heteronemerteans—for which 16S rRNA, COI, 18S rRNA, 28S rRNA, histone H3 genes are available in public databases—shows that Dushia wijnhoffae sp. nov. and Dushia nigra species complex comb. nov. form a clade, which is closely related to Gorgonorhynchus albocinctus Kajihara, 2015 and an undetermined heteronemertean that has been misidentified as Cerebratulus leucopsis (Coe, 1901). Members of Dushia thus show a vicariant Caribbean–West-Pacific distribution, indicating that the lineage predates the rise of the Isthmus of Panama.
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