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Molecular Phylogeny of Echiuran Worms (Phylum: Annelida) Reveals Evolutionary Pattern of Feeding Mode and Sexual Dimorphism

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Molecular Phylogeny of Echiuran Worms (Phylum: Annelida) Reveals Evolutionary Pattern of Feeding Mode and Sexual Dimorphism Molecular Phylogeny of Echiuran Worms (Phylum: Annelida) Reveals Evolutionary Pattern of Feeding Mode and Sexual Dimorphism Ryutaro Goto1,2*, Tomoko Okamoto2, Hiroshi Ishikawa3, Yoichi Hamamura4, Makoto Kato2 1 Department of Marine Ecosystem Dynamics, Atmosphere and Ocean Research Institute, The University of Tokyo, Kashiwa, Chiba, Japan, 2 Graduate School of Human and Environmental Studies, Kyoto University, Kyoto, Japan, 3 Uwajima, Ehime, Japan, 4 Kure, Hiroshima, Japan Abstract The Echiura, or spoon worms, are a group of marine worms, most of which live in burrows in soft sediments. This annelid- like animal group was once considered as a separate phylum because of the absence of segmentation, although recent molecular analyses have placed it within the annelids. In this study, we elucidate the interfamily relationships of echiuran worms and their evolutionary pattern of feeding mode and sexual dimorphism, by performing molecular phylogenetic analyses using four genes (18S, 28S, H3, and COI) of representatives of all extant echiuran families. Our results suggest that Echiura is monophyletic and comprises two unexpected groups: [Echiuridae+Urechidae+Thalassematidae] and [Bone- lliidae+Ikedidae]. This grouping agrees with the presence/absence of marked sexual dimorphism involving dwarf males and the paired/non-paired configuration of the gonoducts (genital sacs). Furthermore, the data supports the sister group relationship of Echiuridae and Urechidae. These two families share the character of having anal chaetae rings around the posterior trunk as a synapomorphy. The analyses also suggest that deposit feeding is a basal feeding mode in echiurans and that filter feeding originated once in the common ancestor of Urechidae. Overall, our results contradict the currently accepted order-level classification, especially in that Echiuroinea is polyphyletic, and provide novel insights into the evolution of echiuran worms. Citation: Goto R, Okamoto T, Ishikawa H, Hamamura Y, Kato M (2013) Molecular Phylogeny of Echiuran Worms (Phylum: Annelida) Reveals Evolutionary Pattern of Feeding Mode and Sexual Dimorphism. PLoS ONE 8(2): e56809. doi:10.1371/journal.pone.0056809 Editor: Donald James Colgan, Australian Museum, Australia Received September 22, 2012; Accepted January 15, 2013; Published February 14, 2013 Copyright: ß 2013 Goto et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Funding: This work was funded by a Japan Society for the Promotion of Science Grant to MK and a Japan Society for the Promotion of Science Research Fellowships for Young Scientists Grant to RG. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript. Competing Interests: The authors have declared that no competing interests exist. * E-mail: [email protected] Introduction In contrast to their systematic position within the annelids, the interfamily relationships of echiurans remain poorly understood Echiurans (spoon worms or innkeeper worms) are a group of [3]. The Echiura has traditionally been classified into the three marine worms with a sausage-shaped non-segmented body and a orders: Echiuroinea (with Echiuridae, Thalassematidae, and highly extensible scoop-like proboscis [1–3]. Approximately 165 Bonelliidae), Xenopneusta (Urechidae), and Heteromyota (with species of echiurans have been described in the shallow to deep Ikedidae) [3]. This classification is based on their morphology, waters of all oceans [4,5]. Most live in burrows in soft sediments, such as the arrangements of the body wall musculature, the although some live in crevices in dead corals or rocks [1–3]. In vascular system, and the hind gut [1–3]. However, this informa- general, they are deposit feeders that collect small organic particles tion is insufficient for understanding the phylogenetic relationships by scooping the sediment with their flattened and elongated among these groups [3]. proboscis, although some are filter feeders that create U-shaped It has also been suggested that the currently accepted burrows and gather food by generating water currents using their classification should be revised [9]. The order Heteromyota was peristaltic bodies [1–3]. originally established for the species Ikeda taenioides (Ikeda, 1904), In recent years, much attention has been paid to the systematic which is endemic to Japan [10]. The body-wall muscle layer of I. position of echiuran worms [6–8]. They were considered to have a taenioides was once described as including outer longitudinal and close phylogenetic relationship with annelids, based on their inner circular layers, which is the opposite of that found in all developmental and morphological characters [1–3]. However, other echiurans [10]. Based on this character, the order echiurans can be distinguished from the other annelids by their Heteromyota was proposed for this species [10]. However, a lack of segmentation [1–3]. Thus, this animal group has been recent detailed assessment of I. taenioides specimens revealed that recognized as a separate phylum [1]. However, recent molecular the arrangements of their muscle layers are not different from the phylogenetic analyses have consistently suggested that echiurans other echiurans [9]. Thus, it was proposed that the order should be placed within the phylum Annelida [6–8]. Thus, they Heteromyota should be abolished and that the family Ikedidae are currently treated as a group of derived annelids [7,8]. should be treated as a junior synonym of the family Echiuridae [9]. PLOS ONE | www.plosone.org 1 February 2013 | Volume 8 | Issue 2 | e56809 Molecular Phylogeny of Echiuran Worms According to this classification, the echiurans comprise the two A molecular phylogenetic analysis of echiurans based on three orders: Echiuroinea and Xenopneusta [9]. genes (16S, 18S, and COI) was recently performed in Lehrke’s It is also interesting that echiuran worms include species with dissertation work [11]. Her results agree with ours in the following and without marked sexual dimorphisms [1–3]. The members of points: (i) Echiura is monophyletic and has a sister group the family Bonelliidae show marked sexual dimorphism [1–3]; relationship with Capitellidae. (ii) The monophyly of Bonelliidae very tiny dwarf males are parasitic on or within the female body. and Ikedidae is highly supported. (iii) The sister group relationship Similar forms of sexual dimorphism were also suggested in the of Echiuridae and Urechidae is highly supported. A high degree of Ikedidae [9]. By contrast, there is no marked sexual dimorphism in congruence with her results supports the conclusion of this study. the families Echiuridae, Thalassematidae, and Urechidae [1–3]. However, there are also some differences between her and our How marked sexual dimorphism evolved in this animal group is results. For example, in our tree (Fig. 2), the monophyly of an intriguing question. Thalassematidae was recovered with strong support (MBP = 94, In order to understand the evolutionary pattern of feeding mode BPP = 1.00), whereas, in her tree [11], it was ambiguous probably and sexual dimorphism in echiuran worms, and to resolve due to the scarcity of the taxa and genes used for the analysis. taxonomic confusion, we constructed the molecular phylogenetic tree of echiuran worms using two nuclear ribosomal genes (18S Evolution of marked sexual dimorphism involving dwarf and 28S rRNA), one nuclear protein gene (histone H3) and one males mitochondrial protein gene (cytochrome c oxidase subunit I, COI). Our results suggested that echiurans are separated into two Representatives of all extant echiuran families were included in the groups; [Thalassematidae+Echiuridae+Urechidae] and [Ikedi- analyses. We discussed the classification and character evolution of dae+Bonelliidae] (Fig. 2). These two groups can be distinguished echiuran worms based on the tree produced. by the presence/absence of marked sexual dimorphism with dwarf males. The members of Echiuridae, Thalassematidae, and Results and Discussion Urechidae do not show marked sexual dimorphism, i.e., the Molecular phylogenetic analysis males and females are similar in size [1–3]. By contrast, those of Bonelliidae and Ikedidae show marked sexual dimorphism, i.e., We collected sequence data for the 18S rRNA, 28S rRNA, H3, very tiny dwarf males, the ciliated bodies of which contain only a and COI genes for the molecular phylogenetic analysis of 15 gonad, a seminal vesicle, and a pair of protonephridia, are echiuran species belonging to nine genera, five families, and three parasitic on or within females [1–3,9]. Our analyses suggested that orders, as well as eight outgroup species (Fig. 1, Table 1, Table Capitellidae is a sister group of Echiura (Fig. 2), as in previous S1). The following outgroups were included to root the echiuran molecular studies [7,8]. The most members of this family are tree; five polychaete annelid species representing three families, sexually monomorphic [12]. Thus, it is highly probable that sexual one sipunculan, and two mollusks (Table 1). The annelid outgroup monomorphism is plesiomorphic for echiurans and that marked included three species of Capitellidae (Table 1), which has been sexual dimorphism evolved in the common ancestor of Bonelliidae suggested as a sister group of Echirua [7,8]. All sequences were and Ikedidae
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