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Barnacle Editor Workshop Barnacle Editor Workshop VLIZ InnovOcean Site Wandelaarkaai 7 – entrance Pakhuis 61 (UNESCO) B-8400 Oostende, Belgium February 24-28, 2020 Final Report Barnacle Participants: Keith Crandall, Meeting Organizer, George Washington University, Washington, DC USA Jens Hoeg, University of Copenhagen, Copenhagen, Denmark Marcos Pérez-Losada, George Washington University, Washington, DC USA Benny Chan, Academia Sinica, Taipei, Taiwan Henrick Glenner, University of Bergen, Bergen, Norway Andy Gale, University of Portsmouth, Portsmouth, United Kingdom Niklas Dreyer, Academia Sinica, Taipei, Taiwan WoRMS Data Management Team (DMT): Stefanie Dekeyzer (Meeting Coordinator) Bart Vanhoorne Wim Decock Leen Vandepitte Target Group: The barnacles – more specifically, the broader group of Thecostraca including the traditional barnacles (Cirripedia) as well as the related groups of Facetotecta and Ascothoracida. The thecostracan barnacles rank among the most commonly encountered marine crustaceans in the world. They deviate from almost all other Crustacea in that only the larvae are free-living, while the adults are permanently sessile and morphologically highly specialized as filter feeders or parasites. In the most recent classifications of the crustacean Maxillopoda 1 and latest phylogenetic analyses 2-4 the Thecostraca sensu Grygier 5, comprising the Facetotecta, Ascothoracida, and Cirripedia, form monophyletic assemblages. Barnacle phylogenetics has advanced greatly over the last 10 years. Nonetheless, the relationships and taxonomic status of some groups within these three infraclasses are still a matter of debate. While the barnacles where the focus of Darwin’s detailed taxonomic work, there has not been a comprehensive review of the species of barnacles as a whole since Darwin. As a consequence, the barnacle entries within the WoRMS Database is woefully out of date taxonomically and missing many, many species and higher taxa. Program: The Thecostraca, which include the Facetotecta, Ascothoracida and Cirripedia, is a highly variable crustacean group in terms of both morphology and biology 6,7. This makes them prime models for studying evolutionary adaptations in both morphology and reproductive systems 8,9. In fact, the specializations in adult morphology, growth, feeding biology and sexual systems prompted Darwin to study cirripedes resulting in one of the first “model organisms” of evolutionary adaptation 10-13. While there has been much recent work on phylogenetic relationships among the barnacles at both the higher 4,14-20 and lower 21-24 taxonomic levels, there has been little integration across levels and across groups working on barnacle evolution. Furthermore, there has not been a recent list of all the species of barnacles since Darwin. We assembled in Ostend at VLIZ a group of the world leaders in barnacle taxonomy, systematics, and evolutionary biology to update the barnacle taxonomy based on the last 20 years of published literature and update and augment the WoRMS database accordingly. We are an experienced group in both taxonomy and phylogenetics with extensive combined experience in producing both web-based and published products of species lists, diversity assessments, phylogenetics, and taxonomy. Our workshop resulted in a number of additions to the WoRMS barnacle taxonomy as well as significant rearrangements reflecting phylogenetic insights into barnacle evolution. Most of our group were new to WoRMS and the workshop also provided training to this outstanding group of barnacle taxonomists to keep the WoRMS taxonomy updated. As dominant and recognizable marine members, the barnacles an exceptional group to target for updating on WoRMS. Activities: Objective 1 – APHIA overview, orientation, anD training. The VLIZ-DMT team led a presentation on the APHIA system, editing, and the barnacle overview. The participants discussed editing standards and targeted efforts for taxonomic updates. Only two in our group had previously edited pages in WoRMS and only one had edited barnacle pages. So this first objective has developed a strong set of barnacle editors moving forward. Objective 2 – Species updates. The Barnacle Group spent the next few days working on species additions for WoRMS. We especially focused on adding a number of fossil taxa, none of which were in the database. Additionally, we added primary literature, diagnoses, fossil ranges, images, and additional information and relevant literature to species pages and added a number of missing species. Objective 3 – Taxonomy manuscript formalize changes. The past 20 years has seen a number of barnacle phylogenetic studies documenting the fact that the current barnacle taxonomy fails to reflect evolutionary relationships. We used the workshop opportunity to develop an updated taxonomy that reflects our current knowledge of the evolutionary history of barnacles (which is substantial). The overarching phylogenetic framework is provided in the manuscript started at the workshop. Our manuscript is now accepted pending revisions at the Zoological Journal of the Linnean Society and will set the standard for barnacle taxonomy for years to come. Our revision is expected to be submitted by the second week of October, 2020. We hope the paper will be published yet in 2020. Objective 4 – WoRMS eDits to reflect upDateD taxonomy. An integral part of our meeting was to not only update the species and higher taxa of missing taxa from WoRMS, but to also update the barnacle taxonomy itself based on the changes formalized in the paper described in Objective 3. To take full advantage of the group of experts being together and to train us in APHIA editing, we made a number of adjustments to the taxonomy to reflect phylogenetic relationships across the barnacles. It was not clear to us that these adjustments should have been made in quarantine until our paper was published. In total, we had hundreds (367) of edits during the week of the workshop and in subsequent weeks (492 the next week, etc.) across the barnacles, again reflecting taxonomy updates (including new species), distributions, images, notes, attributes (feeding type for parasitic barnacles), etc. Because we were making these changes as part of the learning process at the workshop, they caused some consternation on the part of some of our colleagues. Unfortunately, they did not contact any of us nor members of the DMT. Instead, they developed a paper discussing issues of the desire for stability in WoRMS. We, of course, agree and quickly quarantined our changes (thanks to the folks at DMT) until the publication of our article. Nevertheless, our changes will provide a much more robust and lasting taxonomic framework based on sound evolutionary relationships as opposed to the current taxonomy. These edits will come out of quarantine once our paper is published and the barnacle team is committed to updating the barnacle taxonomy in the face of future research. Conclusions: This was an outstanding workshop for those involved in that many of us have been yearning for a comprehensive taxonomy of the barnacles our entire careers. Finally, because of this workshop, we have such a resource. More importantly, this taxonomy is captured in the WoRMS database for easy augmentation in the future. We have trained the next generation of barnacle workers on the WoRMS resource and are dedicated to keeping in up to date. We thank the DMT for their training, insight, and patience with us! Cheers! References 1 Martin, J. W. & Davis, G. E. An updated classification of the Recent Crustacea. Natural History Museum of Los Angeles County Science Series 39, 1-124 (2001). 2 Pérez-Losada, M. et al. The tempo and mode of barnacle evolution. Mol Phylogenet Evol 46, 328-346 (2008). 3 Perez-Losada, M., Hoeg, J. T. & Crandall, K. A. Remarkable convergent evolution in specialized parasitic Thecostraca (Crustacea). BMC Biol 7, 15, doi:1741-7007-7-15 [pii] 10.1186/1741-7007-7-15 (2009). 4 Glenner, H. & Hebsgaard, M. B. Phylogeny and evolution of life history strategies of the parasitic barnacles (Crustacea, Cirripedia, Rhizocephala). Mol. Phylogenet. Evol. 41, 528- 538 (2006). 5 Grygier, M. J. New records, external and internal anatomy, and systematic position of Hansen’s y-larvae (Crustacea: Maxillopoda: Facetotecta). Sarsia 72, 261–278 (1987). 6 Høeg, J. T. & Møller, O. S. When similar beginnings leads to different ends: Constraints and diversity in cirripede larval development. Invertebr. Reprod. Dev. 49, 125-142 (2006). 7 Ruppert, E. R., Fox, R. S. & Barnes, R. D. Invertebrate zoology: A functional evolutionary approach. (Cengage Learning, 2003). 8 Charnov, E. L. in Barnacle biology Vol. 5 Crustacean Issues (ed A. J. Southward) 89-103 (A.A. Balkema, 1987). 9 Høeg, J. T., Pérez-Losada, M., Glenner, H., Kolbasov, G. & Crandall, K. A. Evolution of morphology, ontogeny and life cylces within the Crustacea Thecostraca. Arthropod Systematics & Phylogeny 67, 199-217 (2009). 10 Darwin, C. A monograph of the fossil Lepadidae or, pedunculated cirripedes of Great Britain., (Paleontological Society, 1851). 11 Darwin, C. A monograph of the sub-class Cirripedia, with figures of all the species. The Lepadidae: Or, pedunculated cirripedes., (Ray Society, 1852). 12 Darwin, C. A monograph of the sub-class Cirripedia, with figures of all the species. The Balanidae (or sessile cirripedes); the Verrucidae, etc., etc., etc. (Ray Society, 1854). 13 Darwin, C. A monograph of the fossil Balanidae and Verrucidae of Great Britain. (Paleontological Society, 1855). 14 Buckeridge, J. S. in New frontiers in barnacle evolution. Vol. 10 Crustacean Issues (eds F. R. Schram & Hoeg J. T.) 225-267 (A.A. Balkema, 1995). 15 Harris, D. J., Maxson, L., Braithwaite, L. F. & Crandall, K. A. Phylogeny of the thoracican barnacles based on 18S rDNA sequences. Journal of Crustacean Biology 20, 393-398 (2000). 16 Korn, O. M. in New frontiers in barnacle evolution Vol. 10 Crustacean Issues (eds F. R. Schram & J. T. Høeg) 87-121 (A. A. Balkema, 1995). 17 Mallatt, J. & Giribet, G. Further use of nearly complete 28S and 18S rRNA genes to classify Ecdysozoa: 37 more arthropods and a kinorhynch. Mol Phylogenet Evol 40, 772- 794 (2006). 18 Pérez-Losada, M. et al. The tempo and mode of barnacle evolution.
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