Second International Workshop: Special Issues in Octocoral Taxonomy
Held in Darwin (AUS), Jan 14-24, 2002 at the Museum and Art Gallery of the Northern Territory
Participants: Stephen Cairns, Washington DC (United States), Catherine McFadden, Claremont (California, United States), Gary Williams, San Francisco (California, United States), Götz B. Reinicke, Stralsund (Germany), Jun-Im Song, Seoul (South Korea), Manfred Grasshoff, Frankfurt/Main (Germany), Odalisca Breedy, San José (Costa Rica), Phil Alderslade, Darwin (Australia), Leen van Ofwegen, Leiden (Netherlands), Carlos Sanchez Ortiz, Townsville (Mexico/Australia).
Following the workshop agenda, a collection of discussions and agreements, ideas and plans, was assembled by the participants (collected and edited by G.B. Reinicke)
Updating the Glossary
The Trilingual Glossary, the major outcome of the first Octocoral Workshop in 1981, had been designed to standardize the use of taxonomic terms from historic literature, often written in German and French, to the English language, thus to facilitate proper, precise usage among octocoral workers. It has since become a standard tool for taxonomists needing to reference specific taxonomic terms. From our discussions it also became apparent that it is used as an introductory tool, often given to students who are beginning to work with octocorals, a different use than the sole remaining author, Manfred Grasshoff, had anticipated.
However, in the 20 years since the first edition, there has been a considerable increase in taxonomic terms from new publications, and some terms in the glossary have proved to be inaccurately defined, inadequately coined or obsolete. Publication of a second edition was generally agreed to be a useful enterprise. We thus took about two days to discuss all of the terms in the first edition, decide which definitions required modification and compile a list of new terms to be included in a second edition.
Some good discussion was sparked by the attempt to define the terms coenenchyme, mesenchyme, mesoderm, polyp, colony, etc., in octocorals. A clarification of these key terms will be drafted by Manfred Grasshoff to be included with the glossary’s second edition introductory chapter about octocorals.
Collation of the new and revised terms will be done by Phil Alderslade. The first edition plate originals, still in the Leiden Museum with Leen van Ofwegen, will be scanned and revised for use in the new edition. Stephen Cairns compiled a list of new terms to be included. The production of new and revised definitions will be shared among the group. No time schedule for publication was set.
Major discussion arose following Odalisca Breedy’s suggestion to include Spanish terms in the second edition, based on her translation of the first edition (done in consultation with a linguist). The major objective of this addition would be to provide Spanish speaking workers and students with an introduction to octocorals in their own language.
Serious concern was voiced that we take care to insist on English as THE language of international science, and taxonomy in particular. It was feared that inclusion of Spanish might open the way for other languages, such as Russian, Japanese, Chinese and Korean to be added (Jun-Im Song also had a complete translation of the Glossary with her), further confusing the use of taxonomic terms.
There was widespread support for publication of a separate (bilingual) Spanish edition. This would be in the format: English terms (Spanish translation) – Spanish definition, with an index of the Spanish terms. However, this translation would need to be supplemented by addition of the new terms for the revised second edition glossary. Odalisca is working towards publication of this manuscript.
The question of whether to include Spanish terms in the second edition of the tri- (or quatri-) -lingual glossary remained undecided at the workshop. The original intention of the glossary’s first edition was to provide a language tool to facilitate standardized communication, and at the time "practical considerations precluded the inclusion of [other languages]…" Some workshop participants felt that the introduction of Spanish and possibly other languages could be a major move in support of this goal and that the international Anglo-American language community is well enough established that the adoption of well-defined Spanish (and Korean) terms carefully related to the English terms would not unduly interfere with international communication. It could also be a significant stimulus to Octocorallia research in Central and Latin American as well as in Asian countries. It is possibly the English speakers who are carelessly neglecting papers in languages we can’t read, such as Russian or Chinese.
Developing greater links with molecular biologists.
Interest in developing greater links with molecular biologists was reflected by inviting C. McFadden, Jun-Im Song and C.S. Ortiz to participate in the workshop. Following a brief introduction to molecular systematic methodology by C. McFadden, they reported their current work on Stereonephthya and Scleronephthya (JIS), and Muricea and Pacifigorgia (CO). Fundamental molecular techniques were explained in general terms, and many questions were answered.
Primary targets for molecular biological research in octocorals.
To date, molecular studies of octocorals have addressed a variety of questions at different systematic levels, generally starting with well-established taxa. They range from population studies carried out at species and genus levels (e.g. Cathy McFadden on Alcyonium, Jun-Im Song on Dendronephthya, Götz Reinicke on Briareum sp.) to broader surveys of family-level systematics and phylogeny) on selected sections of Alcyonacea (e.g. Berntson et al., 2001). Several mitochondrial genes have been identified recently that promise to provide reasonable phylogenetic resolution at the genus and family levels and, several groups of researchers are currently using them to construct molecular phylogenies for particular octocoral groups. The use of nuclear intron sequences for low-level phylogenetic studies is also being explored.
Gary Williams and Phil Alderslade expressed their interest in molecular studies of the scleraxonian groups where classical soft corals merge into classical gorgonians. A general conclusion was that there are many problem areas that could be addressed using molecular approaches, but not yet enough molecular systematists working on octocorals.
Berntson EA, Bayer FM, McArthur AG & France SC 2001: Phylogenetic relationships within the Octocorallia (Cnidaria: Anthozoa) based on nuclear 18S rRNA sequences. Mar. Biol. 138: 235-246.
Intraspecific and geographic variability - how to proceed.
Allozymes are still the most effective molecular tool for assessing intraspecific variability and species boundaries in octocorals, and for evaluating the taxonomic significance of morphological variation. Unfortunately, this technique requires fresh or frozen material and thus cannot be applied to museum specimens. The extreme lack of variability in the cnidarian mitochondrial genome has meant that DNA sequencing approaches that are useful for phylogeographic studies in other animal groups have not been applicable to octocorals. This discussion did not reach any distinct conclusions, except to express hope that perhaps the development of new molecular tools may help us in the future.
Sub-ordinal levels in Alcyonacea.
We considered that input from molecular phylogenetics would greatly assist us to better understand sub-ordinal relationships within the Alcyonacea. Molecular studies that have addressed this problem to date have generally used genes that have not been variable enough to resolve sub-ordinal relationships with confidence. Use of more variable genes, such as some of the recently identified mitochondrial regions, will hopefully begin to shed light on sub-ordinal groups.
Octocoral Internet sites
Gary Williams presented his "Octocoral Research Centre (ORC)" at the California Academy of Sciences site: www.calacademy.org/research/izg/orc_home.html.
[… writes G. Williams in Dec. 2001:] "The site is still in a rather [preliminary] and unfinished form, with much formatting still to be done, but has (or will have) the following ten headings:
As an immediate result of the workshop, the Sea Pen Bibliography has been linked with Dr. F.M. Bayer's Bibliography of Octocorallia web site, formerly based at the Smithsonian Institution (see below). The workshop participants further suggested setting up separate (or linked) bibliographies of molecular systematic work on octocorals including cladograms, and chemical literature (e.g. secondary metabolites etc.). The Tree-of-Life octocoral site will also eventually be linked.
At present the ORC appears to be the most comprehensive Octocoral Internet site available. Gary offered to let the site become the major forum for exchange of information concerning octocorals (see also next section). Participants were invited to submit details about themselves, such as their research interests and availability to help people.
Keeping Bayer’s Octocoral bibliography site up to date
We discussed transferring the existing bibliography of Prof. Frederic Bayer at http://nmnhgoph.si.edu/gopher-menus/Octocorals.html to the Octocoral Research Centre at the California Academy of Science, to be maintained by Gary Williams and his staff. Dr. Bayer subsequently agreed to allow the site to be moved there. It has successfully been transferred as of 15 May 2002. Workshop participants and other octocoral workers will contribute to updating the bibliography (1996 to present). Subsequent collection and cataloguing of Octocoral papers will occur on a weekly basis.
Ted Bayer’s crystallography data and unfinished papers.
Stephen Cairns partly changed his field of work and joined retired Dr. F.M. Bayer to inherit bits of his octocoral knowledge and experience and to assist in sorting out his tremendous archives, including manuscripts, SEM pictures and other results of his lifetime work. Stephen thus will be the reference contact for requests concerning Dr. Bayer’s work and will assist him in carrying on the various threads of his research. Stephen can also be contacted for reprint requests.
The task of compiling a World list of nominal Octocoral taxa.
We agreed to use Leen van Ofwegen’s (Alcyonacea) and Gary Williams' (Pennatulacea) lists compiled in 1995 for the UNESCO register of marine species as starting points for a state-of-knowledge world list. Octocoral taxa will be divided up among workshop participants who will check and revise the list. Subsequently (in March ‘02) Leen sent the current version of the Alcyonacea list to participants and solicited expressions of interest from individuals willing to take responsibility for particular taxa. A start has been made with an update on April 20, 2002.
Recruiting research workers to our discipline to continue after we retire.
In planning the workshop a number of requests from younger students wishing to participate were rejected in order to keep the group at a size that would facilitate meaningful discussion. It was felt that the emphasis of the workshop should be on detailed discussion of general problems in the field rather than "teaching".
One conclusion of our discussions was that we need to advertise to attract students to the field, and the best medium for that in the long term may be through the Cal Academy web site. Recruitment of young colleagues will be a major goal of the octocoral symposium to be held during ICCB 2003 (see below). A Young Workers Forum could be established to explicitly encourage students to present their work for discussion, comments and support. Jun-Im Song reported the lack of jobs in taxonomic fields for the students she trains in South Korea. Financial support needs to be found and organized to allow young taxonomists to enter the profession once they have completed their training (this issue obviously is one of concern to taxonomists in all fields).
Running beginners training courses Octocoral taxonomy.
Jun-Im Song reported that in Seoul students from various backgrounds have expressed interest in octocoral taxonomy, and there is thus a need for an integrated training program for university students. Recently, some museums in South Korea have started to award PhD degrees, and many universities have joined the PEET program for training graduate students and post docs. Particular jobs including taxonomic expertise need to be established to attract students to the field.
Phil Alderslade reported on his experiences running taxonomy training workshops in Phuket (Thailand) and Suva (Fiji). Many students often attend these courses, but most of them are not planning careers in taxonomy – they attend for other reasons. Again, those that may think about a career in taxonomy have trouble getting jobs. "South Pacific island nations may provide a surprising source of taxonomists in the future as they have much aid funding to set up new positions whereas the developed nations are not funding an increase in museum type departments and positions." [PA]
Literature access in less privileged countries.
Discussion of this question followed comments presented by J.I. Song and O. Breedy. The solution that is usually suggested is to contact experts directly and ask them to send literature. This approach does not always succeed because those experts (often) don’t have (or don’t take) the time to respond. Literature can usually be obtained during direct visits to the appropriate institutions (e.g. literature of F. Bayer and S. Cairns through the Smithsonian Institution). But requesting copies of old literature and papers via mail in many cases appears to be impractical and regularly fails. Typical responses are that papers are too long, no staff is available to photocopy them and there are limited (or no) funds to mail them.
So what to do? We did not reach any decision about establishing a central reference center or system, and there seems to be little we can do about it at this time, except provide support to foreign workers when approached directly. The octocoral bibliography on the ORC website and large scientific library databases can serve as sources for references and access to the respective publications, and the ORC website will further assist communication among researchers. Access will certainly improve as more journals make their archives available electronically and internet access extends in other countries.
An international octocoral conference
It was suggested that we organize an Octocoral mini-symposium
to be held during the upcoming 6th International Conference on Coelenterate
Biology (ICCB), scheduled in Kansas (US) in 2003. Manfred Grasshoff contacted
Daphne Fautin, the conference organizer, and she enthusiastically agreed to
include an extra octocoral section in the conference (which still needs to be
further conceptualized and communicated). Subjects suggested for the symposium
included:
- The life of Willy Kükenthal, or other octocoral specialists of the past;
- Expedition Reports;
- Young Workers Forum.
Regular specialist forums.
This subject was discussed in conjunction with scheduling an octocoral symposium as part of ICCB in 2003. We considered such meetings to be generally useful, and it was suggested that we hold another workshop in 5 years. Stephen Cairns offered to host it in Washington DC in 2007.
Standardising the style of our research papers.
The question of a standardized format for octocoral taxonomic papers was discussed, specifically including a quick reference section (single "tear-out" page) at the end of the paper that could be photocopied to put in a binder for that group of octocorals. A couple of arguments in favor were countered by the problems of general international implementation and having to conform to the different styles and publication formats of various journals. Nevertheless, Phil Alderslade agreed to draft a summary, quick reference page for people to look at in a paper in preparation for "The Beagle – Records of the Museum and Art Galleries of the Northern Territory".
Sclerites and Biometry.
Generally, it appeared to the participants that biometric analysis of sclerites and colony forms is not a very valuable pursuit for taxonomic purposes. Sclerites can be measured by confocal microscopy and digital image analysis used to differentiate types. But they vary intraspecifically with geography, depth, in color and life history. Different branching patterns and growth forms in colonies of Gorgonacea, and form and size variations in Alcyonacea also depending on the state of extension and contraction of preserved material in particular further obscure these characters as taxonomic tools [JIS].
The current state of Xeniidae taxonomy.
Taxonomy of Xeniidae genera has been helped by Phil Alderslade’s recent description of six new genera, based mainly on characters of the sclerite microstructure. Phil has an on-going project to investigate sclerite structure in all nominal Xeniidae genera. At the species level, however, revision of the larger (Xenia, Heteroxenia) and less common genera (Cespitularia) is needed. Such an effort would require further re-collections from type locations and SE Asian reefs. Molecular techniques should be applied wherever possible.
An inventory of unstudied museum collections.
Various collections and material from different (far-off) locations sit on museum shelves awaiting study and publication. To further facilitate and support comprehensive biogeographic studies and comparison, Goetz Reinicke suggested assembling a list of collection material that is available for study. The list will be prepared and started by Goetz, and will include information about whether material is identified and catalogued, and the predominant taxa in the collection. It will then be made available through Gary Williams’ ORC site (see above).
The compilation of type inventory lists from various museums would facilitate requests for reference material from colleagues. Museum type inventories are increasingly becoming available as databases on the Internet, and are sometimes searchable like that of the Smithsonian NMNH collection. But until more are available, direct correspondence with museums obviously remains the standard option for locating type material.
Leen van Ofwegen’s (Alcyonacea) and Gary Williams' (Pennatulacea) projected World List of Nominal Octocoral Taxa also contains fields for the type status and could optionally be supplemented with information on museum, collection numbers and type localities. But this will require long-term work.
Stephen Cairns suggested that reviving the old practice of depositing isotypes in other collections would increase accessibility to (type) specimens in museums. These could further be supplemented by slides with sclerites and (desirably, if available) voucher specimens for/from molecular studies.
Other contributions to the Workshop
Goetz Reinicke gave a presentation introducing the German Oceanographic Museum in Stralsund, highlighting their long tradition of successful exhibition work as well as fields of scientific specialization: Coral reef ecology, soft coral taxonomy, fish ecology and marine mammals.
Manfred Grasshoff gave an introduction to the "Frankfurt Theory of Evolution" – engineering morphology and the reconstruction of the main evolutionary lineages of animals. The lecture focused on the evolution of coelenterates. The octocoral’s typical coenenchyme-structure appears to be relatively close to that of the (hypothetical) ancestral coelenterates. This forms the basis for the evolution of the various so-called Hexacorallia and fossil Tetracorallia, while the Tetrazoa (= Medusozoa) evolved independently. The latter appear to be highly derived, and none of the groups (Hydrozoa, Scyphozoa, Cubozoa) can be referred to as the basal group of coelenterates. Molecular studies yield similar results, with the Tetrazoa being characterised by a specific molecular feature (linear mitochondrial DNA) that is not ancestral (Bridge et al. 1992, Grasshoff & Gudo 2002).
Bridge D, Cunningham CW, Schierwater B, De Salle R, Buss LW 1992: Class level relationships in the phylum cnidaria - evidence from mitochondrial genome structure. Proc. Nat. Acad. Sci. US 89: 8750-53.
Grasshoff M, Gudo M 2002: The origin of metazoa and the main evolutionary lineages of the animal kingdom – the gallertoid hypothesis in the light of modern research. Senckenb. leth. 82: 307-326.
Acknowledgements
Cordial thanks are due to Phil Alderslade and his colleagues, who invited the workshop to the Museum and Art Gallery of the Northern Territory and carefully provided all the extensive infrastructure necessary for concentrated and productive working atmosphere. "Thanks" to all the MAGNT’s staff – nice days for fishing anyway. Further thanks go to the workshop participants, who contributed to this manuscript, and to Cathy McFadden for brushing up the linguistics.
Research interest of participants:
Jun-Im Song (South Korea)
(1) the species diversity of soft corals, genus Dendronephthya living in the temperate zone nearby the western Pacific Ocean, as my morphological study after the phylogenetic analysis of the genus based on molecular makers. (2) the coevolution of zooxanthellae associated with anthozoan taxa using molecular and morphological studies since three years. As my side activities, I am trying to set up both ways for the next generation, the establishment of a Korean PEET program and new Natural History Museums.
Götz B. Reinicke (Germany)
Ongoing studies on soft corals collected during expeditions to the Chagos Archipelago and Socotra (both Indian Ocean) are pursued in collaboration with Leen van Ofwegen (Leiden, NL). Further collections of Xeniidae from SE Asian reefs are studied to revise old collections and descriptions, with particular regard to the genera Xenia, Heteroxenia and Cespitularia. These should preferably be supplemented by population genetic data of those species. A small study on morpho-functional constraints of sclerite characters variation is proceeding on the genus Briareum.
Catherine S. McFadden (United States)
Ongoing studies of the molecular systematics of soft corals, in particular the construction of a molecular phylogeny to elucidate relationships among genera in the family Alcyoniidae. Molecular evidence for patterns of reticulate evolution among European Alcyonium species. Past research focused on population genetic structure and life history evolution in the genus Alcyonium, and demographic and population genetic consequences of clonal reproduction in an alcyoniid from the NE Pacific.