PORCUPINE MARINE NATURAL HISTORY SOCIETY

NEWSLETTER

April 2002 Number 10

25th anniversary

25th anniversary

ISSN 1466-0369

Porcupine Marine Natural History Society

Newsletter

No. 10 April 2002

Hon. Treasurer MEMBERSHIP Jon Moore Ti Cara Chairman Point Lane Julia Nunn Cosheston Cherry Cottage Pembroke Dock 11 Ballyhaft Rd Pembrokeshire Newtonards SA72 4UN Co. Down BT 22 2AW 01646 687946 [email protected] [email protected] Porcupine MNHS welcomes new members- scientists, students, divers, naturalists and Hon. Editor lay people. We are an informal society Frances Dipper interested in marine natural history and 7 Rutland Green recording particularly in the North Atlantic Hilton and 'Porcupine Bight'. Members receive 3 Huntingdon newsletters a year which include Cambs proceedings from scientific meetings. PE28 9NT 01480 830507 Individual £10 Student £5 [email protected]

COUNCIL MEMBERS

Mike Bailey [email protected] Ian Killeen [email protected] Roger Bamber [email protected] Paul Brazier [email protected] Peter Barfield [email protected] Anne Bunker [email protected] Jon Moore [email protected] Susan Chambers [email protected] Julia Nunn [email protected] Frances Dipper [email protected] Ivor Rees [email protected] Frank Evans [email protected] Shelagh Smith [email protected] Judy Foster-Smith [email protected] Peter Tinsley [email protected] Annette Little [email protected]

CONTENTS

Editorial 1-2

Council and AGM Minutes 3-5

Accounts 6

Porcupine meetings, Other meetings 7

Book Reviews 8

PORCUPINE PIECES First British Record of a European barracuda Sphyraena sphyraena . Doug Herdson 9

A D’Arsy Thompson transformation of crotchets to uncini and its implications for the phylogeny of polychaetes. Peter Gibson 10-11

PAPERS FROM PORCUPINE 2002: Changes in Marine Biology over the last 25 years.

The Chinese mitten crab, Eriocheir sinensis: the introduction of an alien species. Roni S. Robbins and Paul F. Clark 12-16

The Firth of Forth Spatial Study – Taxonomic problems associated with long-term monitoring. Susan Hamilton 17-22

Meiofauna in marine pollution studies and environmental assessments. Stephen Jarvis 23-26

Long-term changes in the zooplankton community of the North Sea. Richardson, A.J., Lindley, J.A. and Edwards, M. 27-32

Challenges in Marine Biodiversity: The Global View (abstract). A. D. McIntyre 33

Marine technology: challenges of the last frontier (abstract). D. Meldrum 33-34

Practical developments in numerical analysis techniques used for marine biological surveys. Annette Woodham 34-39

EDITORIAL Address to PORCUPINE on the The observant amongst you will notice occasion of its 25th Anniversary. that this issue of our Newsletter has a different cover to normal! This David Heppell continues the celebration of Porcupine’s 25th anniversary, begun at our AGM and annual meeting in As one of the Founding Fathers of Edinburgh over 14th-16th March 2002. PORCUPINE, and its first Hon. The meeting was a great success both Treasurer, I very much regret I cannot in social and scientific terms and our be with you all today to share in sincere thanks must go to Susan the celebration of PORCUPINE's 25th Chambers who organised the meeting Anniversary. As many of you know I and the National Museums of Scotland was taken seriously ill last August and who hosted it. If any of you are in any for a while was not expected to doubt about just how much work is live. However, I did survive and am involved in organising such an event, very grateful for all the prayers try it yourselves! 78 names appear in and e-mails and cards expressing your the delegates list and the majority of good wishes for my recovery. I am those did actually attend. This may be writing this from my wheelchair in the a Porcupine record. Eight of the hospital in Vancouver where I am papers presented are written up in this learning to walk again. issue either in full manuscript (6) or as abstracts (2). Further papers will follow This is meant to be merely a in the November issue. congratulatory address, and not a history of the Society, although I hope The conference dinner followed the someone else will write up a summary first day’s proceedings and was held at of PORCUPINE's first 25 years before the Edinburgh zoo, courtesy of the they fade from our collective memory. Edinburgh Royal Zoological Society – Back in 1977 I was involved in very appropriate for Porcupines?! It discussions about launching a new was a strange experience wandering marine biological Society which would through the zoo at night with no-one have its emphasis on the fauna and except the animals and fellow flora of the North Atlantic and Porcupines for company. The Mediterranean (my own area of excellent meal and bar were much interest) and marine recording in appreciated. particular (the main interest of our Founding Mother, Shelagh Smith). I The Saturday morning saw an intrepid was keen to honour the memory of band meeting at 8.30 am at Dunbar to John Gwyn Jeffreys, author of "British investigate the rocky shore there. The Conchology" and participant in the weather was kind and the tide exploratory dredging cruises of HMS sufficiently low to allow several hours "Lightning" and HMS "Porcupine" in of ferreting around and recording. A 1868-1870. The success of these full report and lists of species recorded cruises led to the subsequent will hopefully be published in the next circumglobal expedition of HMS newsletter and interesting finds added "Challenger". Rather than call the to the Recording Scheme records. Society the Jeffreys Society, which would have placed too much emphasis Reproduced below is an address on malacology (as Jeffreys was one of kindly sent by David Heppell, our first the great British conchologists), Honorary Treasurer, currently living in we decided on PORCUPINE, after the Canada. ship and, in the early years, always spelled all in capitals to distinguish it from the lower case mammal. For the same reason it was not called

PMNHS Newsletter No. 10 April 2002 1

"The Porcupine Society" as we as well as the academic wanted to encourage the inevitable traditions of PORCUPINE for the next question "What is PORCUPINE?" and 25 years and beyond. thereby possibly gain a new recruit.

From the beginning the Society held IMPORTANT NOTICE! both field meetings and indoor Our Honorary Treasurer, Jon Moore meetings, combined wherever and his wife Ginny have produced a possible. As so many meetings of potential new Porcupine member – a interest to marine biologists were little girl called Kate. Congratulations always held in London, we wanted to to them both! meet on the fringes. Early meetings were held in Orkney, South Shields, Guernsey, Portaferry, Cardiff, Menai, Plymouth, Millport and the Isle of Man, with themes ranging from marine parasites to meiofauna. Although many members were malacologists we soon had members interested in algae, sponges, polychaetes, crustaceans, pycnogonids and fish. The benefit of the field meetings was that we all learned from each other, sharing our knowledge and experience of collecting techniques and methods of identification. No longer need one write "Lithothamnion sp.' SUBSCRIPTION REMINDER or "Chiton sp." into the records! Could all members please note that annual subscritions are now (over)due The Newsletter was a great success for 2002 - £10 for full members, £5 for from the beginning, produced then students. If you are unsure if you have from Gestaetner wax stencils, under paid (many of you pay by bankers the editorship of Fred Woodward. order) please e-mail Jon Moore on Later Frank Evans introduced the [email protected] delightful little cartoons of marine Porcupines. We even had a Porcupine mascot at one time - where is it now? We decided our Society did not need a President - no figurehead, only working Officers. Shelagh Smith COPY DEADLINES Extended to July 1st for the next was our first Hon. Secretary and st Dave McKay the Records Convener - issue and October 1 for the the idea being to provide a vehicle November issue for recording those groups for which there was no other Recording Scheme in operation. This was one of the early ideas that did not take off, but I think in all other ways the Society has held true to its original aims and ambitions.

I wish the Society continuing good www.pmnhs.org.uk fortune, and trust that the present day Porcupines will carry on the social

PMNHS Newsletter No. 10 April 2002 2

Summary of Minutes of the Council Action: FD to e-mail constitution to all Meeting held on March 14th 2002 at Council members and publish it in the the National Museums of Scotland, next newsletter. Edinburgh. 4. The accounts for the year ended Present: Julia Nunn, Frank Evans, 31st December 2001 were presented Ivor Rees, Shelagh Smith, Roger by the Hon Treasurer, Jon Moore. Bamber, Susan Chambers, Annette Little, Frances Dipper, Judy Foster- 5. A report on membership was given Smith, Jon Moore, Anne Bunker. by Jon Moore. He reported that Appologies: Bridget Betts membership was stable, with 17 new members this year. 1.The Edinburgh Conference was Action: Membership drive to be reported to be running smoothly. discussed at the autumn Council Council members were pleased with meeting. the turnout, the venue and the organisation of the Conference. 6. A report on the PORCUPINE Council expressed its thanks to Susan Newsletter was given by the Hon Chambers. A problem had occurred editor, Frances Dipper. She requested with some speakers expressing copy for the next issue and ideas for surprise at being asked to pay the topics. She also suggested a special conference registration fee. It was cover for the 25th anniversary edition. decided that in future, invited speakers It was decided to try the idea of a should not be asked to pay a different habitat theme for each edition registration fee, but would be asked if (related to the Habitat Action Plans?) the organisation they work for (in the and to have a silver cover for the case of speakers who are not self anniversary edition (No. 10 April employed) would like to contribute to 2002). All Council members agreed PORCUPINE by donating the fee. that the book reviews were a good Action: Susan Chambers to write to idea and should be continued, with the speakers and refund registration fees addition of a list of newly published for this conference. books. Action: FD to approach publisher It was suggested that PORCUPINE concerning silver cover. could seek sponsorship in future if it looked as if the conference budget 7. Jon Moore reported that some would not stretch to this. records had been received for the Action: To be discussed at the Recording Scheme. autumn council meeting. Action: JM to compile list of members e-mail addresses. 2. Council members. Nigel Grist – dropped from Council. 8. Anne Bunker reported on the web Peter Tinsley – to continue as a site and asked for a policy on use by Council member. non-members. It was decided that Bridget Betts – resigned. non-members wanting to put Ivor Rees – would like to resign when information requests on the web site a suitable replacement has been should be encouraged to join but that found. one request was acceptable for non There are currently 2 vacancies. members. Council decided to ask Peter Barfield Action: AB to encourage non- and Paul Brazier if they would like to members making requests to join become Council members. PORCUPINE.

3. All Council members need to read 9. Frank Evans reported that progress the constitution. on the poster was ongoing.

PMNHS Newsletter No. 10 April 2002 3

members paying their subscriptions by 10. It was decided that the 2002 field standing order. Over the year the cost meeting would be in N. Wales with the of the production of the newsletter has details to be confirmed. benefited from the use of a cheaper but equally good printer. The meeting 11. The venue and organiser(s) for the at Brampton, Huntingdon showed a 2003 conference. welcome profit. Society membership Action: Jon Moore to approach Andy stood at 194 of whom 164 were Mackie at the National Museum of ordinary members. This year we have Wales. enrolled 17 new members, a welcome increase. 12. The History of PORCUPINE is ongoing. All information should be The report was accepted following passed to Shelagh Smith. proposal by Roger Bamber, seconded by Ralph Robson.

Hon. Editor, Frances Dipper MINUTES OF THE 25TH ANNUAL There have been three issues of GENERAL MEETING OF Porcupine Newsletter this year. PORCUPINE MARINE NATURAL However, only about half of those who HISTORY SOCIETY gave papers at the Huntingdon meeting have submitted reports for Held at the National Museums of publication. Within the newsletter the Scotland, Edinburgh on Thursday 14th sections headed Porcupine Pieces March 2002 (in the same room as the and Porcupine Problems have become inaugural meeting of 1977). well established. Porcupine Pieces carries articles and reports submitted Chairman: Julia Nunn by members while Porcupine Problems deals with information requests and replies. The latter ran to 1. Apologies for absence between two and four items per issue There were no apologies for absence. this year. The success of this section was illustrated by the topic of mantis 2. Minutes of last AGM shrimps, which contributors kept going These had been published in the for three issues of the newsletter. March 2001 issue of Porcupine Reports from Plymouth and the Isle of Newsletter and were accepted. Purbeck field meetings are still pending. 3. Matters arising from the minutes of the last AGM The report was accepted following There were no matters arising from the proposal by Annette Little, seconded minutes. by Anne Bunker.

4. Officers' reports. Chairman, Julia Nunn. Hon. Treasurer, Jon Moore There have been two Council The unaudited Treasurer's report was meetings, at Carlisle in November and presented to the meeting (Editor’s in Edinburgh during the current annual note: the audited accounts are meeting. Grateful thanks were offered included in this issue of Porcupine to Mike Bailey and Frances Dipper for Newsletter). The Society's funds were their work in organising the annual in a satisfactory condition. The meeting in March 2001 at the Treasurer would like to find a Environment Agency in Huntingdon, repository giving a better return for the and Annette Little and Anne Bunker Society's deposit but this would were thanked for their labours with inevitably make difficulties for those registration. The Chairman further

PMNHS Newsletter No. 10 April 2002 4 thanked Susan Chambers and her and this was agreed with no team for their very successful objections. organisation of the current meeting. Officers for the next year: The report was accepted following Hon. Chairman, Julia Nunn proposal by Frank Evans, seconded Hon. Treasurer, and Hon. Records by Judy Foster-Smith. Convenor Jon Moore Hon. Editor, Frances Dipper 5. Website (www.pmnhs.org.uk). Anne Bunker reported that the website now lists details of the meetings of the Council members for the next year: Society. A table of contents of the Mike Bailey, Roger Bamber, Anne current Porcupine Newsletter appears, Bunker, Peter Barfield, Paul Brazier, together with a print-out of "Porcupine Susan Chambers, Frank Evans, Judy Pieces" from its pages. Our standard Foster-Smith, Annette Little, Ian recording form is reproduced, details Killeen, Ivor Rees, Shelagh Smith, of how to become a Society member Peter Tinsley are shown and there is a "Features" item, changed from time to time. Input 9. Other business from Porcupine members to this last There being no other business the was requested by the webmaster. Chairman declared the meeting closed. 6. Poster. Frank Evans reported that the delays in producing an advertising poster have now been overcome and the work, newly undertaken by Joshua Arnott of Whitley Bay, is advancing well.

7. Field meetings. The Chairman expressed the thanks of the Society to the organisers of the Dorset field meeting. It is proposed that the next field meeting will be at a location in Anglesey in September 2002.

8. Election of Officers and Council members. No proposal was made to elect a secretary. The Chairman stated that if desired all the other Officers were willing to continue in post, while Bridget Betts and Nigel Grist did not wish to stand for re-election to Council. Peter Barfield was proposed for Council membership by Susan Chambers, seconded by Annette Little and Paul Brazier was proposed for Council membership by Ivor Rees, seconded by Christine Howson. There being no other nominations the chairman proposed that Officers and Council members be elected en bloc

PMNHS Newsletter No. 10 April 2002 5

PORCUPINE RECEIPTS AND PAYMENTS ACCOUNT for the year ended 31 December 2001 (unaudited)

Year to 31.12.00 Year to 31.12.01 £ £ £ £

RECEIPTS 223 Subscriptions- 1997 - 1999 10 1572 2000 15 40 2001 1376 20 2002 5 1855 1406 76 Bank Interest (net of tax) 69 10 Sale of PN Back Number 14

1941 Total Receipts 1489

PAYMENTS 1965 Newsletter- Printing 476 438 Postage 321

2403 Total Newsletter Costs 797 5 Bank charges 0 357 Membership leaflets - printing 0 94 Recording Scheme cards - printing 0 183 Council meeting expenses (travel/catering) 163 0 Repayment of overpaid subscriptions 24 3042 984

(1101) SURPLUS (DEFICIT) BEFORE MEETINGS 505

361 MEETINGS – Plymouth (2000) (99) MEETINGS – Huntingdon (2001) 278

(839) SURPLUS (DEFICIT) FOR THE YEAR 783

4208 BALANCE BROUGHT FORWARD 3369

BALANCE CARRIED FORWARD 1503 Current Account 747 1866 Deposit Account 3404 3369 4151

Jon Moore, Hon Treasurer 10th March 2002

PMNHS Newsletter No. 10 April 2002 6

MEETINGS, MEETINGS, MEETINGS, MEETINGS, MEETINGS

PORCUPINE MEETINGS 25th June 2002. Adapting to Climate Change: Overcoming the barriers to change. CoastNET, London : Progamme details on www.coastms.co.uk PORCUPINE FIELD MEETING 2002 8-12th July 2002. Estuarine and September 2002 in Anglesey. Details Lagoon Fish and Fisheries. will be e-mailed/sent to all Porcupine Fisheries Society of British Isles members when available. Annual International Symposium. University of Hull. www.hull.ac.uk/iecs

THOUGHTS ECSA Seaweed Workshop. Either If anyone would like to volunteer to organise either the annual scientific March/April or late summer 2002. meeting or a field meeting for future Contact Martin Wilkinson: 0131 451 years, please contact Julia Nunn. 3468, [email protected]

Similarly if you have ideas about where you would like such meetings held and their content, again please contact Julia or any council member.

OTHER MEETINGS

22nd May 2002. Making Waves: Tackling Litter in the Aquatic Environment - National Aquatic Litter Group NALG London, Keynote speaker Michael Meacher For programme go to www.coastms.co.uk

13-15th June 2002. Info’Coast 2. 2nd European Symposium on Knowledge and Information for the Coastal Zone. Noordwijkerhout, The Netherlands. (postponed from October 2001). [email protected] +44 223 667050.

19-20th June 2002. Coastal Futures Scotland 2002 Edinburgh. Coastal Management for Sustainability. Full programme details on www.coastms.co.uk

PMNHS Newsletter No. 10 April 2002 7

procreation. It makes you realise how vital diversity is and just how much we REVIEWS should value it.

The author (our editor, no less!) should be congratulated on the way she has skilfully presented such a huge amount of information in such a small book. She has brought together anecdotal observations and sound scientific insight and distilled them in a clear and yet tantalising way, carefully explaining the more complex issues. She has chosen excellent examples to illustrate her points and, as a true conservationist, has taken the opportunity to touch on the continuing need for the stewardship of our marine Extraordinary Fish and freshwater systems. By Frances Dipper (BBC Worldwide Ltd. 2001) My only real (but very minor) criticism is the unfortunate over-use of different Review by Judy Foster-Smith font types and sizes. I have yet to work out why some bits of the text This glossy little paperback is part of a have been singled out for special font series produced as a follow up to ‘The treatment when their relevance Blue Planet’ programmes which, I’m appears much the same as the rest. sure, need no introduction for All they do is interrupt the reader’s ‘Porcupines’. The book is a credit to flow. the BBC. It is a delightful synthesis of the lives of some of the weirdest However, the book is colourfully animals in our waters, aimed at the illustrated with lots of exotic informed public. photographs and will certainly appeal to anyone interested in wildlife, the From the bioluminescent powers of the sea, and …. extreme survival. At £7.99 deep-sea angler fish to the wildly it’s a gem. Order one now at abstract markings of the Picasso www.amazon.com! triggerfish, and from the ugly, warty frogfish, which doesn’t swim at all, to the graceful Sargassum flying fish which not only swims, but also takes to the sky when pursued by predators, the book is full of fascinating facts.

I felt it a treat to read, like a day out, being taken on a whistle-stop tour of the amazing array of loony-looking fish and the bizarre ways in which they are adapted to survive and feed in extreme conditions. Then on to a show of sounds and light and body language – the art of fish communication, and, finally, to a display of the most elaborate means of underwater

PMNHS Newsletter No. 10 April 2002 8

and lower flanks which are diagnostic PORCUPINE PIECES of S. barracuda.

This specimen appears to be a European Barracuda Sphyraena sphyraena on the basis of the size, elongated body shape, fully-scaled pre-operculum, dark elongated head (like a rocket's nose cone with the upper jaw fitting neatly into a protruding lower one) and approximately twenty dark vertical half bars along its body.

The European Barracuda Sphyraena First British Record of a European sphyraena, is called a Great barracuda Sphyraena sphyraena Barracuda in a number of books, but

this name should kept for the Douglas Herdson Caribbean/West Atlantic Sphyraena National Marine Aquarium, Plymouth barracuda. The European Barracuda

th can grow to 160 cm and 38 kg. (The On the 25 November 2001 the crew record for Sphyraena barracuda is of F.V. Regina Maris, skipper David 200cm and 50 kg.) Kessel, was fishing for hake with a monofilament set net when they netted The fish was on display in ice at the a barracuda. They were fishing 8.5 Quayside Fish Centre in Porthleven, miles south east of the Lizard, and is now frozen until Mr Strike Cornwall. The fish was sent to Newlyn decides whether to have it stuffed or Fish Market where it was bought for sent up to the Natural History Museum £16 by John Strike of the Quayside in London. Fish Centre in Porthleven, Cornwall.

A single unusual fish like this is not a The fish is 106 cm (total length) and sign of global warming, but it may fit 4.2 kg (gutted, so probably an into a pattern when correlated with ungutted weight of over 4.5 kg), and is numbers of fish records as we are the first record of a barracuda in British starting to do at the National Marine waters. Aquarium.

The fish was examined by Dr Paul Photo by Phil Monckton Gainey and photos were sent to me and Alwynne Wheeler (formerly of the

Natural History Museum) and we have all agreed on the identity of the fish.

Of the three species found in the North

East Atlantic and Mediterranean only

Sphyraena sphyraena extends north to the Atlantic coast of the Iberian

Peninsula and southern Bay of Biscay, also neither of the other species exceeds 65 cm TL. The western

Atlantic Sphyraena barracuda would be an outside possibility, but this fish lacks the pale tips to the lobes of the tail and the black blotches on the belly

PMNHS Newsletter No. 10 April 2002 9

A D’Arsy Thompson transformation needed. In the present study a of crotchets to uncini and its diagram of a crotchet from the spionid, implications for the phylogeny of Aonides paucibranchiata, (used in the polychaetes previous study) was drawn freehand on a grid of squares. It was then Peter Gibson replotted on a grid of straight lines which were cut by curves - the ICAPB, University of Edinburgh, West distances between these were Mains Road, Edinburgh, UK progressively doubled (see figure). The crotchet was, as a result, D’Arsy Thompson in his classic book transformed into an uncinus. More On growth and form* showed how to convincing transformations might be transform the fish Diadon to the achieved using other dimensions for sunfish Orthagoriscus. The method the second grid. Here, I am simply points to, amongst other things, the attempting to illustrate the principle. importance of allometric growth Allometric growth would achieve the (relative changes in proportions of same result. Indeed, the shape of structures) in evolution. In biological chaetae probably depends upon terms, transformations become varying growth rates for linked irrelevant once a group of organisms macromolecules within the structures. or structures are known to be related Families with crotchets or uncini live to one another - this is the essence of in, or on, a variety of substrata (see evolutionary homology. For example, it table) and their chaetae appear to is easy to see how four classes of show appropriate structural echinoderms evolved from a crinoid adaptations. Crotchets are probably ancestor without knowing what the adapted for digging burrows: they selection pressures were. The have a long shaft for leverage, a systematics of the polychaetes is a far subterminal projection which may act greater problem; there are large as a pick and frequently a crest of number of families and no easy way of teeth probably for gripping the sides of connecting their phylogenies as has the tube. The crotchets of species been achieved with vertebrates. Dales living in soft substrata are frequently in his book Annelids (Hutchinson hooded and these hoods probably act University Library, 1962) used the as scoops. The uncini, in contrast, are feeding structures to help show short and always bear numerous relationships between polychaete teeth; each has an asymmetrically groups (Fig. 9, p. 75). Fauchald and placed basal ligament and muscle to Jumars looked at feeding guilds in pull the chaeta into the body to detail but did not discuss phylogeny release it from gripping the sides of the (Oceanography and Marine Biology tube; they act as ratchets. Annual Review, 1978). The transformations described When considering methods for suggest that species with crotchets characterising chaetae (Gibson, gave rise to species with uncini. The Robson & Armitage, Newsletter of the crotchets are likely to have been Porcupine Marine Natural History derived from simple spines (acicular Society No. 2, 1999) I felt there might chaetae) which are frequently seen be some relatively simple within crotchet bearing species. An mathematical definition for some argument for the reverse direction is chaetae and that transformations difficult to make when one considers could also be used to convert complex other structures and in particular those images to simpler ones as well as used in feeding (see Dales). The saying something about phylogeny. transformation may have a similar For example, an ellipse may be application to other types of chaetae. described by
r2 if one knows what transformation (mapping function) is

PMNHS Newsletter No. 10 April 2002 10

I wish to thank Dr. Cosens for checking the manuscript for shaky English and silly mistakes.

*A simple account is given by Peter and Jean Medawar in Aristotle to Zoos (Oxford University Press, 1985) under Form and mathematics (p. 102) and Transformations (p. 264).

Table. A selection of sedentary polychaete families bearing crotchets or uncini and the relationship between chaetae to the habitat (Fauvel’s polychaete keys and Fauchald & Jumars).

Crotchets Uncini Family Substratum/tube Family Substratum/tube Cirratulids* Soft, hard/variable Ampharetids Soft/mucus Spionids Soft, hard/variable Terebellids Soft/mucus Capitellids Soft/variable Pectinarids Soft/sand Arenicolids Soft/temporary Sabellids** Soft/sand Maldanids Soft/mucus Serpulids Surface of/limy Chaetopterids Soft/horny Sabellariids Surface/sand Plus spines * & crotchets**

Figure 1. D’Arsy Thompson transformation of a crotchet to an uncinus, a) freehand drawing of the crotchet of Aonides paucibranchiata plotted on a grid of squares and b) the result of replotting the crotchet on a grid with equally spaced radiating straight lines, and curved lines where the distance between them is progressively doubled.

PMNHS Newsletter No. 10 April 2002 11

PORCUPINE 2002. CHANGES IN MARINE BIOLOGY OVER THE LAST 25 YEARS

Papers from the PMNHS meeting held at the National Museums of th Scotland, Edinburgh from 14 -16th March 2001

In the UK, the mitten crab has been reported from the Humber, Medway, Tyne and Thames catchments. The establishment of this species within these river systems has potential environmental implications nationally. The risk of further invasions along the eastern counties may be considered low but is probably dependent on the The Chinese mitten crab, level of shipping traffic and coastal Eriocheir sinensis: the dispersal of larvae from estuaries introduction of an alien where the mitten crab is abundant. species

Introduction Roni S. Robbins and Paul F. Clark The Chinese mitten crab, Eriocheir sinensis H. Milne Edwards, 1854, is a The Natural History Museum, Cromwell native of east Asia, with a distribution Road, London SW7 5BD, England from Fukien Province, China, ca. 26ºN, to the western areas of the Synopsis Korean Peninsula ca. 40ºN. The life Eriocheir sinensis, the Chinese mitten history of the Chinese mitten crab is crab, is a native of China and the atypical for crabs, spending most of its western coastal regions of Korea. It life in fresh water, but returning to the predominantly lives in freshwater but sea to reproduce. Males and females migrates seawards in order to migrate downstream during late reproduce. In 1912 this alien crab was summer and attain sexual maturity in first recorded from Europe. A estuaries. It is during these mass specimen was captured from the River migrations that the crabs are most Aller, a tributary of the Weser, visible. After mating the females are Germany and the species was thought to continue seaward into probably introduced into this river via deeper water to overwinter before the discharge of ballast water from a returning to brackish water in the commercial vessel. This invasive crab spring to hatch their eggs (Panning, has now spread throughout north-east 1939). Larval development probably Europe with a reported distribution occurs in the lower estuary, with from Finland to Portugal including juvenile crabs gradually migrating England. Its recent appearance and upstream into fresh water to complete subsequent establishment in San the life cycle. These upstream Francisco Bay, USA, is also thought to migratory movements can be be from the discharge of ballast water. substantial with juvenile mitten crabs migrating up to ~ 1500 km in their

PMNHS Newsletter No. 10 April 2002 12 native habitat whilst growing to adult and Belgium this non-native crab has size. been reported in pest proportions.

Additional aspects of the mitten crab North America life history of interest include its The mitten crab has also been burrowing behaviour, which has the reported from North America with potential to accelerate riverbank records from the Detroit River at erosion. Furthermore the Chinese Windsor Ontario and Lake Erie, mitten crab is regarded as a delicacy Canada (Nepszy & Leach, 1973), the in the Far East especially while Mississippi Delta (Horwath, 1989) and sexually mature. However, E. sinensis San Francisco Bay (Cohen & Carlton, is the second intermediate host of the 1997), United States. However the oriental lung fluke, Paragonimus only region where the crab appears to westermanii (Kerbert, 1878): if the have established a reproducing crab is eaten raw or is poorly cooked, population is San Francisco Bay. The the parasite can infect humans, American wildlife authorities are so causing the disease paragonimiasis. concerned about the spread of this alien species and its burrowing Since 1912, mitten crabs have behaviour that live mitten crab imports become widely dispersed from their into California in 1987, and native Asian habitat by being subsequently the whole of the United inadvertently transported to Europe States in 1989, have been banned by and recently into North America, via legislation (see Horwath, 1989). the ballast water of ships. These invasions of E. sinensis have the Distribution in England potential to cause a number of The Chinese mitten crab was first environmental problems. recorded from the British Isles when a specimen was captured on the intake European distribution screens of the Lots Road Power The first record of Eriocheir sinensis in Station, Chelsea in 1935 (Anon, Europe was in 1912 when a single 1935a,b; Harold, 1936). There were no male was captured from the River further reports until 1949 when a Aller, a tributary of the Weser, second specimen was reported from Germany (Panning, 1939). Its present Southfields Reservoir, near Castleford estimated distribution ranges from in Yorkshire (Cockerham, 1949) and Finland (Haahtela, 1963) in the north, the Humber (Clark, 1984). through Sweden, Russia, Poland, Denmark, Germany, the Czech There was a lack of mitten crab Republic, the Netherlands, Belgium records from the Thames until the mid- and England to France. The 1970s when Ingle & Andrews (1976) southernmost Atlantic coast record is reported capturing this species at in the Golfe de Gascogne, France West Thurrock Power Station. After its (Vigneux et al., 1993), but the crab has establishment, the Thames mitten crab extended its range via the Garonne population remained at relatively low canal system to Sigean, Languedoc- and constant numbers through the late Roussillon, a Mediterranean district of 1970s and 1980s. southern France (Petit, 1960). Recently this crab was reported from Thames update the Tagus Estuary, Portugal (Cabral & To update distribution data in the Costa, 1999), but the extent of this Thames catchment, The Natural invasion is to date unknown. History Museum, supported by an Populations in the northern part of the Environment Agency grant, made an Baltic and the Sigean lagoon system appeal to the general public for mitten have not become established, crab sightings. This information was whereas in Germany, the Netherlands plotted onto a map of the Thames

PMNHS Newsletter No. 10 April 2002 13 catchment area by Clark et al. (1998 Austropotamobius pallipes figure 2). The Chinese mitten crab is (Lereboullet, 1858), which is already now known from the west as far as the under considerable threat from foreign River Colne at Staines and is present crayfish introduced into British rivers. in most of the Thames tributaries A further concern is that E. sinensis is downstream of this point. In the east, a burrower (Panning, 1939), and a the crab has been found in the rivers mass invasion of all tributaries east of Cray, Darent, Quaggy, Pool and Staines could result in significant Ravensbourne. In the north-east, riverbank erosion. Burrowing could sightings are common in the rivers threaten unprotected engineering Roding and Lee. The most northerly earthworks. This concern was one of report of E. sinensis is from the River the contributing factors that resulted in Lee tributary at Enfield, some 15km the banning of live mitten crab imports upstream off the River Thames. The into the United States. Burrowing survey also recorded E. sinensis in behaviour has been confirmed in the every tributary from Chelsea upstream grounds of Syon Park, Middlesex. to Chertsey. These include the Beverley Brook and the Rivers Coarse fishing has also reportedly Wandle, Brent, Duke of been affected in the Thames Northumberland, Crane, Hogsmill, catchment. Anglers along the River Longford, Ember, Mole and Ash. The Lee and at a private club near the furthest upstream record at was River Cray record that during the Staines, ca. 65 km from Tilbury. migration period during late summer, crab strikes on bait virtually prevent Environment Agency data analysed sport fishing. from power station collections at West Thurrock (1976–1993) and Tilbury Recent UK records (1993–1996) also indicated that the Recent UK records include Kingsnorth Thames mitten crab population is now Power Station, the Medway, 1990 firmly established and, since 1992, has (NHM reference collection); small been increasing in numbers as well as numbers from the River Tyne, 1998 geographical range. These records (Ken Watson, pers. comm.); a female show that small annual numbers of crab from the mouth of the Teign mitten crabs were collected at West Devon, 1999 (Martin Attrill, pers. Thurrock Power Station in the late comm.) and Dungeness, Kent, 1970s and 1980s. However the mitten 2001(NHM reference collection). crab population increased in 1992 and that trend continued in 1993 until the Control of alien mitten crabs power station closed in March. That Prevention is the best method of this increase has persisted, is controlling the invasion of exotic indicated from numbers at Tilbury, species into new regions and this from 1993 to 1996. requires international co-operation. One of the main methods of Thames fallout transportation of species around the The reported distances that mitten world is in ballast water of shipping crabs can migrate in China and (Carlton, 1985). Recent attempts to Europe suggest that the whole of the use Open Ocean Exchange or Thames river system is accessible to freshwater flushing of ship’s ballast invasion and dispersal via canal may not be appropriate methods to systems (Petit, 1960) linking control bio-invasions (see Hülsmann & catchments, is a reality. Further Galil, 2001 for discussion). population expansion could eventually threaten freshwater habitats and In the specific case of mitten crabs, communities including those currently extensive trapping at a particular point occupied by the native crayfish is an option. Not only is this method

PMNHS Newsletter No. 10 April 2002 14 labour intensive, but Peters & Hoppe Andrews, M.J., Aston, K.F.A. Rickard, (1938) attempted to control mitten crab D.G. & Steel, J.E.C., (1982). The populations in Germany by a similar macrofauna of the Thames Estuary. method during the migration season The London Naturalist, 61: 30–61. without success. There is, however, a Cabral, H.N. and Costa M.J., (1999). potential for commercial exploitation On the occurrence of the Chinese because the mitten crabs are regarded mitten crab, Eriocheir sinensis, in as a gastronomic delicacy. Portugal (Decapoda, Brachyura). Restaurants in Hong Kong and Japan Crustaceana, 72(1): 55–58. charge a high price for such a meal. Mitten crabs may fetch up to £5.55 per Carlton, J.T., (1985). Transoceanic pound in London when in season. An and interoceanic dispersal of coastal added attraction of exploiting the marine organisms. Marine Biology European mitten crabs population for Annual Review, 23: 313–371. food is that the establishment of parasitic lung disease is thought Clark, P.F., (1984). Recent records of unlikely. Firstly, the populations in alien crabs in Britain. Naturalist. Hull. Europe were probably established by London. 109(970): 111–112. plankton i.e. larvae and juvenile specimens transported in ballast Clark P.F., P.S. Rainbow, R.S. water. Consequently the adult Robbins, B. Smith, W.E. Yeomans, M. population would probably be parasite Thomas & Dobson G., (1998). The free. Secondly, Paragonimus Alien Chinese mitten crab, Eriocheir westermanii is specific to a primary sinensis (H. Milne Edwards, 1854) intermediate host of aquatic snails [Crustacea: Decapoda: Brachyura], in assigned to the Thiaridae, and the the Thames Catchment. J Mar. Biol. climate in Northern Europe may be too Ass. 78(4): 1215–1221. cold for members of this gastropod family. However before the Cockerham, H., (1949). Crab in commercial export of mitten crab can reservoir. Angling News, 2090: 414. be sanctioned, tests for parasites should be undertaken. Cohen, A.N. & Carlton, J.T., (1997). Transoceanic Transport Mechanisms: www.nhm.ac.uk/zoology/crab Introduction of the Chinese Mitten Crab, Eriocheir sinensis, to California. References Pacific Science, 51: 1–11. Anon, (1935a). Exhibitions and notices. October 22nd, 1935. Sir Arthur Haahtela, I., (1963). Some new Smith Woodward LL.D., F.R.S., Vice- observations and remarks on the President, in the chair. Proc. Zool. occurrence of the Mitten Crab, Soc. London, 1935: 948. Eriocheir sinensis Milne Edwards (Crustacea, Decapoda), in Finland. Anon, (1935b). Abstracts of the Aquilo Societas Amicorum Naturae Proceedings of the Zoological Society Oulensis, 1: 9–16. of London. October 22nd, 1935. Sir Arthur Smith Woodward LL.D., F.R.S., Harold, C.H.H., (1936). Thirtieth Vice-President, in the chair. Proc. Annual Report on the Results of the Zool. Soc. London, 1935(no. 387): 2. Chemical and Bacteriological Examination of the London Waters for Adema, J.P.H.M., (1991). Krabben van the 12 months ending 31 December Nederland en Belgie. [Crabs from 1935. Metropolitan Water Board Holland and Belgium]. Leiden: London, p. 20-21. Nationaal Natuurhistorisch Museum. Horwath, J.L., (1989). Final Rule on importation of injurious wildlife: mitten

PMNHS Newsletter No. 10 April 2002 15 crabs. Federal Register; Rules and und der angrenzenden Meeresteile. Regulations, 54: 22286–22289. Jena: Verlag Gustav Fischer.

Hülsmann, N. & Galil, B.S., (2001). Vigneux, E., Keith, P. & Noël, P. The effects of freshwater on marine (eds)., (1993). Atlas préliminaire des heterotrophic protists – implications for Crustacés Décapodes d’eau douce de ballast water management. Mar. Poll. France. Coll. Patrimoines Naturels. 14, Bull, 42(no. 11): 1082–1086. Paris: Secrétariat de la Faune et de la Flore, Laboratoire de Biologie des Ingle, R.W. & Andrews, M.J., (1976). Invertébrés Marins et Maalacologie Chinese mitten crab reappears in Muséum National d’Histoire Naturelle, Britain. Nature, London, 263: 638. Conseil Supérieur de la Pêche, Ministère de l’Environment. pp i-vi, 1– Kerbert, C., (1878). Zur Trematoden- 56. Kenntnis. Zoologischer Anzeiger, Liepzig, 1: 271–273.

Lereboullet, A., (1858). Description de deux nouvelles espèces d’écrevisse de nos rivières. Mémoires de la Société des Sciences, Agriculture et Arts de Strasbourg, 5: 1–11.

Milne Edwards, H., (1854). Notes sur quelques Crustacés nouveaux ou peu connus conservés dans la collection du Muséum d’Histoire Naturelle. Archives du Muséum d’Histoire naturelle (Paris), 7: 145–192.

Nepszy, S.J. & Leach, H.J., (1973). First records of the Chinese mitten crab, Eriocheir sinensis (Crustacea: Brachyura) from North America. Journal of the Fisheries Research Board of Canada, 30: 1909–1910.

Panning, A., (1939). The Chinese Mitten Crab. Report of the Board of Regents of the Smithsonian Institution (Washington), [1938] 3508: 361–375.

Peters, N. & Hoppe, H., (1938). Uber bekampfung und verwertung der Wollhandkrabbe. Mitteilungen Hamburgischen Zoologischen Museum und Institut, 47: 140–171.

Petit, G., (1960). Le crabe chinois est parvenu en Méditerranée. Vie et Milieu, 11: 133–136.

Schellenberg, A., (1928). Krebstiere oder Crustacea. II. Decapoda, Zehnfüßer. Die Tierwelt Deutschlands

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the sub tidal benthos and sediments of The Firth of Forth Spatial the estuary had not been Study – Taxonomic problems systematically studied until then. associated with long-term monitoring The Firth of Forth Benthic Faunal Survey Susan Hamilton The first Firth of Forth Benthic Faunal Centre for Marine Biodiversity and Survey was carried out in 1976 - 1977 Biotechnology, Department of Biological by Heriot-Watt University, in Sciences, Heriot-Watt University, collaboration with the then Forth River Riccarton, Edinburgh, EH14 4AS, Purification Board, and funded by the Scotland. Manpower Services Commission. 73

sample stations were chosen, covering Introduction the Firth from the Forth Bridges to the The Firth of Forth is a drowned river Isle of May, two van Veen Grab valley forming an embayment of the samples were taken at each station North Sea. Along its shores lie many and the benthic infauna identified and industrial developments and enumerated. The findings, together residential areas, including Edinburgh, with results of a similar survey of the Scotland’s capital city. Historically, its upper Forth Estuary, were published waters have received a mix of by Elliot and Kingston (1987). agricultural and urban run-off, treated and untreated sewage effluent, The Forth Spatial Study discharges from shipping and many Much has changed since that first other man-made pollutants, and, by survey was carried out in 1976. A the 1970s, had become one of the suite of legislation has been most intensively used sea-areas introduced to improve and protect the around Scotland. marine environment preceding the

recent implementation of the Urban Despite much of the coastline and Wastewater Treatment Directive and inter-tidal area being designated as a driving investment in increased protected area in one way or another sewage treatment. The result is a in order to conserve wetland habitats significant improvement in water and the internationally significant quality. It was decided to investigate numbers of bird life that they support, the effects of these improvements on

PMNHS Newsletter No. 10 April 2002 17

the benthic environment and their Faune de France 1923 and 1927, animal communities by conducting a Hartmann – Schröder’s repeat survey, the Forth Spatial Study, Borstenwurmer, 1971 and Day’s in 2000, almost 25 years after the Polychaeta of Southern Africa, didn’t original, using the same sampling always agree. However, assuming stations and strategy as before to these species were correctly identified allow direct comparisons to be made. in both surveys this was not a problem The present study is a collaborative – new names could simply be venture between SEPA, Scottish substituted for old. Natural Heritage, Heriot-Watt University and the National Museums However, there was also a of Scotland, being funded by them as considerable change in the number of well as by East of Scotland Water and taxa recorded overall, in the numbers Shell U.K. of taxa found at many stations, and of their identities. Ignoring indeterminate and juvenile categories, there seemed Polychaete faunal trends 1976-2000 to be an increase in the total number So what changes have occurred in the of taxa found from 135 to 161, (~20%), polychaete fauna of the Firth of Forth however only half of those found in over these 25 years? The workup of 2000 were also recorded in 1976. The the samples is still ongoing, we only number of taxa found at the majority of have polychaete data from 48 of the stations had also risen. 71 stations sampled so far, but when this was tabulated alongside the full The greatest increases were seen in 1976 dataset, difficulties were the inner Firth, while the decreases encountered arising from considerable seem to be mainly located in an area differences in nomenclature and to the east of the island of Inchkeith. taxonomic resolution Firstly, 30 As yet we have not found any obvious species have simply had their name explanation for this, but these results changed since 1976. Now, for the have yet to be correlated with those from the other

Percentage Change in Number of Polychaete Taxa 1976 - 2000 component parts of the 300 study. As for the stations showing

200 substantial increases in numbers of taxa, where 100 have all these extra species come from? Has there been 0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 21 22 23 24 25 26 27 29 30 31 32 33 34 35 36 37 38 39 40 42 50 53 55 60 62 66 71 72 a dramatic Percentage Change improvement in the diversity of

-100 the fauna or Stations have some or even all of sake of consistency, we can use the these perceived changes arisen from Species Directory, (Howson & Picton advances in our taxonomic knowledge 1997), to standardise nomenclature, and in the literature now available, but then there was no recognised enabling us to distinguish between standard and the key texts, Fauvel’s more species? A few groups of

PMNHS Newsletter No. 10 April 2002 18

Polychaeta are selected as examples material guaranteeing a degree of to illustrate this effect. consistency in their identification.

Aphroditoidea Pholoidae The only taxa recorded in both years Everything was identified as Pholoe were relatively large, robust and minuta in 1976 – at least the ‘minuta’ distinctive - Aphrodita aculeata, in Fauvel and Hartman-Schröder were Gattyana cirrosa and Lepidonotus both the same, however neither are squamatus. Gattyana amondseni was the true ‘minuta’ as described by recorded in 1976, since cited in the Fabricius in 1780. Now we have a lot literature, however in retrospect this more species in the literature to chose may well have been either G. cirrosa from, with even more confusion as variation in this species has been between the different texts. noted (Tebble & Chambers 1982), or perhaps Harmothoe antilopes – these Pholoe ‘inornata’ - as ‘minuta’ species are similar to the untrained (Howson & Picton, 1997), in eye. Two specimens of H. antilopes Chambers 1985, and Chambers & were recorded in 2000, although not at Muir 1997, but as ‘baltica’ in Petersen same station as G. amondseni in 1998 1976. Unfortunately, the faunal Pholoe ‘inornata’ - Petersen 1998, material from 1976 was burned in ash- similar to but distinct from free dry weigh biomass ‘synopthalmica’ determinations, only a few specimens Pholoe 'baltica' - remaining from outlying reference Pholoe ‘tuberculata’ - Southern stations. One important lesson 1914, may be valid species learned from that survey therefore, (A.S.Y.Mackie), similar to ‘baltica’ was the importance of keeping Pholoe’ synophthalmica’ - similar to properly collated reference collections! but distinct from ‘inornata’ (Petersen, In 1976, species of Harmothoe 1998), valid species (Howson & recorded were extenuata, imbricata Picton, 1997), Chambers 1985, and and lunulata, while species recorded in Chambers & Muir 1997 2000 were impar, pagenstecheri, Pholoe ‘assimilis’ - Petersen 1998 andreapolis, marphysae, arenicolae Pholoe pallida - The only distinctive, (also including lunulata?) and eyeless species, agreed on by antilopes. This observed change is Chambers 1985, Petersen 1998, and probably due to much improved Chambers & Muir 1997and also descriptions, keys, including Tebble Christie 1982 as cf. Anoculata and Chambers (1982) and Chambers and Muir (1997), in the Linnean Three species of Pholoe were Society Synopsis series, and their recorded in the Forth this time – P. illustrations - some of Fauvel’s were pallida, P. ‘inornata’ and P. quite fanciful. The perception of what ‘synopthalmica’, the only species constituted key anatomical features included in the new Linnean Society differed between Fauvel and Guide (Chambers & Muir 1997). Hartmann-Schröder, as did their However, as there is at least one other synonymies. There is still much work distinct species occurring in the central to be done on this group, especially in North Sea which keys out to re-evaluating older collections. Also, Petersen’s ‘assimilis’, this group still there is the question of observer needs some clarification. It is certain, experience, as, when one first starts however, that all Pholoe species out looking at these animals they can occurring in the Forth would have all look remarkably similar, only years been lumped together as ‘minuta’ in spent looking at thousands of 1976. specimens and keeping reference

PMNHS Newsletter No. 10 April 2002 19

Nephtyidae In Hayward & Ryland 1990 it states While Nephtys caeca, N. hombergii, N. that “probably only one species (of incisa, and N. longosetosa were Magelona) occurs in northern recorded in both years, N. hystricis European waters.” The new Species was recorded only in 1976, and N. Directory (Howson & Picton 1987) kersivalensis and N. assimilis only in names 5 species – M. alleni (including 2000. From this it appears that 2 new M. rosea), M. filiformis, M. minuta, M. species were found in 2000 and one mirabilis and M. wilsoni, and Fiege, was lost. However, from several Licher & Mackie (2000) later papers published by Rainer, (1984, distinguished a sixth, M. johnstoni, 1989, 1990, 1991) we know that previously included in M. mirabilis. In historically the descriptions of N. 2000, Magelona alleni, filiformis, hystricis and N. incisa have been minuta, and johnstoni, were all found, confused, and that N. kersivalensis raising the number of species from 2 and N. assimilis have been re- or 3 to 4, purely owing to increased described and elevated to full species taxonomic resolution. status from being varieties of N. hombergii. Again some material from Cirratulidae 1976 fortunately survived, was re- While 7 taxa were recorded in 1976 - examined and could be reassigned. Caulleriella caput-esocis, Caulleriella As expected, N. hystricis was re- sp. (assumed to be different from C. identified as N. incisa, N. incisa proved caput-esocis), Chaetozone setosa, to be small specimens of N. Cirriformia tentaculata, Dodecaceria kersivalensis and, although no N. concharum, Tharyx (now hombergii were re-examined, they Aphelochaeta) marioni, and Tharyx probably included some N. assimilis. (now Aphelochaeta) multibranchiis So although a change in the fauna there were 10 in 2000 - Caulleriella appears to have taken place since alata, Caulleriella zetlandica, 1976, this is in fact not the case, the Chaetozone setosa, Chaetozone taxonomy has simply moved on. gibber, Cirratulus cirratus, Cirratulus caudatus, Cirriformia tentaculata, Magelonidae Dodecaceria sp., Aphelochaeta Fauvel included only 2 species, M. marioni, and Tharyx killariensis. papillicornis now synonymised with Only 4 taxa were recorded in both ‘mirabilis’ and having distinctive years - Cirriformia tentaculata, inflated ends to some setae on the 9th Aphelochaeta marioni, (the segment, and M. rosea, described as identifications of both these species similar to papillicornis but with no were confirmed from surviving 1976 mention of any pigmentation. specimens), Chaetozone setosa, in all Hartman-Schröder included 3 species, it’s myriad guises and Dodecaceria sp. M. alleni – a large species with a and D. concharum, assuming they are broad pigmented band, M. papillicornis the same animal. (mirabilis) and M. minuta – a small slender species with 1 tooth over the Although Caulleriella caput-esocis was main fang of its abdominal hooks. In described by both Hartmann-Schröder 1976, three taxa were recorded, and Fauvel, our 1976 Forth specimens Magelona alleni, with the broad were probably either Chaetozone pigmented band, Magelona rosea, and gibber, which was only described in Magelona sp. These last two were 1994 (Woodham and Chambers 1994) both common, but as they weren’t and/or Caulleriella zetlandica, another recorded at the same stations they large species equally and relatively should maybe both be lumped under numerous and difficult to distinguish Magelona spp. between from incomplete or damaged material. 1 specimen remaining from 1976 was re-identified as C. ?gibber

PMNHS Newsletter No. 10 April 2002 20 but this was in such poor condition it what may be more a reflection of an was impossible to be sure. The increase in our knowledge and identities of Tharyx multibranchiis and experience of these fascinating Caulleriella spp? remain uncertain. animals over the last 25 years.

The situation has much improved References since James Blake’s partial revision of Blake, J.A. (1991). Revision of some Tharyx, Aphelochaeta and Monticellina genera and species of Cirratulidae in 1991, Unicomarine’s 1996 review of from the western north Atlantic. the family, and Woodham and Ophelia Suppl. 5, 17-30 Chambers attempts to resolve some of the problems within Chaetozone, but Chambers, S.J. (1985). Polychaetes much still needs to be done. This is from Scottish Waters. Families another group where observer Aphroditidae, Sigalionidae and experience definitely contributes to the Polyodontidae. Royal Scottish ease and certainty of correct Museum Studies 2, 38pp identification. I certainly would not like to attempt to identify Cirratulidae now Chambers, S.J. & Muir, A.I. (1997). using the literature available in 1976. Polychaetes: British Chrysopetaloidea, Pisionoidea and Aphroditoidea. Summary Synopsis of the British Fauna No. 54. Of course, there are many other key papers published since 1976 that Christie, G.(1982). The reproductive contribute to the increased number of cycles of two species of Pholoe off the species likely to be recorded around Northumberland coast. Sarsia, 67 (4), our coasts. These include Hartmann- 283-292 Schröder’s revised Annelida, Borstenwurmer (1996), Frederik Day, J.H. (1967). A Monograph on the Pleijel’s review of the Phyllodocidae Polychaeta of Southern Africa. Part 1. (1989) and Torleif Holthe’s Polychaeta Errantia. Trustees of the British Terebellomorpha (1986), as well as Museum (Natural History) London. 1- Brendan O’Connor’s work on the 458 Glyceriidae (1987), John Hartley’s 1981 paper exploding the myth of the Day, J.H. (1967). A Monograph on the ubiquitous Aricidea jeffreysii, and Polychaeta of Southern Africa. Part 2. others too numerous to mention here Sedentaria. Trustees of the British but gratefully acknowledged by Museum (Natural History) London. anyone faced with a dish of worms to 459-878 identify. And we must not forget the important part we all play ourselves by Elliot, M. and Kingston, P.F. (1987). recording and sharing our The Macrobenthos of the Estuary and observations and specimens at Firth of Forth. Proceedings of the workshops and seminars, and in this Royal Society of Edinburgh, Series B newsletter. 93 pts 3 & 4, 449-465.

So, as far as the Firth of Forth Spatial Fauvel, P. (1923). Polychetes Study is concerned, until all the data is Errantes. Faune De France. Paris.5. collected, species reassigned where 488pp possible, or lumped where not, it is impossible to say what proportion of Fauvel, P. (1927). Polychaetes the perceived change in the species Sedentaires. Faune De France. composition suggested by these Paris.16. 494pp preliminary results might represent a real increase in the diversity of the Fiege, D., Licher, F. and Mackie, polychaeta in the Firth of Forth, and A.S.Y. (2000). A partial review of the

PMNHS Newsletter No. 10 April 2002 21

European Magelonidae: Magelona Rainer, S.F. (1984). Nephtys pente sp. mirabilis redefined and M. johnstoni n. and a key to Nephtys from Northern sp. nov. distinguished. J. Mar. Biol. Europe. J. Mar. Biol. Ass. U.K. 64, Ass. U.K. 80, 215-234 899-907

Hartley, J.P. (1981). The family Rainer, S.F. (1989). Redescription of Paraonidae in British waters: a new Nephtys assimilis and N. kersivaliensis species and new records with a key to and a key to Nephtys from Northern species. J. Mar. Biol. Ass. U.K. 61, Europe. J. Mar. Biol. Ass. U.K. 69, 133-149 875-889

Hartmann-Schröder, G, (1971). Rainer, S.F. (1990). The genus Annelida, Borstenwürmer, Polychaeta. Nephtys in northern Europe: Die Tierwelt Deutschlands. Veb redescription of N. hystrix and N. Gustav Fischer Verlag Jena. 58, incisa. Journal of Natural History. 24, 594pp 361-372

Hartmann-Schröder, G, (1996) Rainer, S.F. (1991). The genus Annelida, Borstenwürmer, Polychaeta. Nephtys of northern Europe: a review 2., neubearbeitete. Auflage. Die of species, including the description of Tierwelt Deutschlands, Veb Gustav N. pulchra sp. n. and a key to the Fischer Verlag Jena. 58 Nephtyidae. Helgoländer Meeresuntersunters. 45, 65-96 Holthe, T. (1976). Polychaeta Terebellomorpha. Marine Southern, R. (1914). A Biological Invertebrates of Scandinavia. 7, 194pp Survey of Clare Island in the County of Mayo. Ireland. Section 2, Hayward, P.J. & Ryland, J.S. (eds). Archiannelida and Polychaeta. 1990. The Marine Fauna of the British Proceedings of the Royal Irish Isles and North–west Europe, 2 Academy. 31. (17 - 47). 1-160. volumes. Clarendon Press, Oxford. Tebble, N. and Chambers, S. (1982). Howson, C.M. & Picton, B.E. (eds). Polychaetes from Scottish Waters. (1997). The Species Directory of the Family Polynoidae. Royal Scottish Marine Fauna and Flora of the British Museum Studies 1, 73pp Isles and surrounding seas. Ulster Museum and the Marine Conservation Unicomarine Ltd. (1996). A Guide to Society, Belfast and Ross-on-Wye, the Family Cirratulidae including a key 508pp. to Anterior Portions. (Unpublished).

O'Connor, B.D.S. (1987). The Woodham, A. and Chambers, S. Glyceridae of the North Atlantic and (1994). A new species of Chaetozone Mediterranean, with descriptions of from Europe, with a re-description of two new species. Journal of Natural Caulleriella zetlandica. Mém. Mus. History. 21, 167-189 natn. Hist. nat. (Actes de la 4eme Conference international des Pleijel, F. (1988.) Phyllodoce from Polychetes). 162, 307-316. Northern Europe. Marine Invertebrates of Scandinavia 8, 159pp

Petersen, M. (1998). Pholoe from Northern Europe: a key and notes on the nearshore species. J. Mar. Biol. Ass. U.K. 78, 1373-1376

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meiofauna is its high diversity. Meiofauna in marine pollution Species rich assemblages of studies and environmental nematodes and copepods are assessments frequently accompanied by the larval stages (termed “temporary meiofauna”) of larger, more familiar marine invertebrates as well as some Stephen Jarvis more obscure meiofaunal groups. Emu Ltd, Hayling Island Marine These include the little–known Laboratory, Ferry Road, Hayling Island, kinorhynchs, gastrotrichs and Hampshire PO11 0DG. gnathostomulids which, although [email protected] widespread, are seldom encountered even by professional zoologists. Introduction Because of their unique features they This paper is not intended as an are ranked as phyla in their own right, exhaustive review of pollution effects and are exclusively meiobenthic. on meiofauna. More comprehensive Tardigrada and Loricifera also fall into treatments can be found in Coull & this category. In all, twenty-two of the Chandler (1992) and Giere (1993) and metazoan phyla have meiobenthic these works should be consulted for representatives. further reading. The purpose of this short contribution is to provide a brief Because these animals are so summary of the reasons for and abundant and diverse they are against their use and to point out that potentially an extremely useful source there are circumstances, more of information for mankind. They are frequent than often thought, where helping to shed light on aspects of high quality samples can be obtained evolution, speciation and relatively easily. Used in conjunction biogeography but, on a more with recent identification literature, immediately practical level, they can meiofauna samples can form the basis also be used to track long- and short- of effective and meaningful term changes in the marine environmental assessments. environment.

What is meiofauna? Why should they be used in Meiofaunal animals are those taxa that environmental assessments? are routinely thrown away in biodiversity studies and environmental Analysis of marine macrofauna has assessments. They are traditionally long been used in ecological studies defined as benthic metazoans and is a standard tool in environmental (including sometimes Foraminifera) assessments and monitoring that pass through a 0.5 mm sieve and programmes where the potential are retained on a 63 (or 42) micron effects of man’s activities in the mesh. They occur in most aquatic oceans are of concern (e.g. oil and habitats from mid-continental streams gas development, aggregate and lakes to the abyssal ocean plains, extraction). Meiofauna is also useful and can reach phenomenal densities, in this respect and arguments for using frequently more than 1 million it as an indicator of environmental individuals per square metre of change have been cogently set out sediment, sometimes an order of elsewhere (most recently by Kennedy magnitude greater. Taxonomically, in & Jacoby, 1999). Three pertinent marine habitats, meiofauna is almost attributes have been touched upon invariably dominated by nematodes, already, namely the sheer abundance, usually followed in number by the ubiquity and diversity of meiofauna. harpacticoid copepods. However, one However, it has two distinct theoretical of the fascinations of studying advantages over most macrobenthic

PMNHS Newsletter No. 10 April 2002 23 taxa. The first relates to rapid be used in toxicity or “effects” testing. generation times. Many nematodes In vitro studies on harpacticoids have and harpacticoids reproduce shown that they are often more frequently and life cycles can be sensitive than nematodes. Easily completed in a matter of weeks. This measurable sub-lethal responses such means that meiofauna communities as clutch size or development time can react more rapidly to environmental be highly informative. These tests perturbations which may then be may then be conducted with species detected in changes in community from the local environment and so attributes such as diversity or provide some predictive modelling “evenness”. This responsiveness is capability relevant to the study area. extremely useful where early warning of environmental degradation is What are the disadvantages? required in sensitive areas, and could There are, of course, disadvantages in be achieved by frequent sampling and using meiofauna in pollution studies. It analysis of the meiofauna. Meiofaunal is certainly true, for instance, that sampling improves temporal resolution much more is known about the of survey data. autecology of macrofaunal species. This makes it possible to make use of The second advantage is that the gross changes in feeding mode of meiobenthic species complete their certain groups (e.g. polychaetes) by whole life cycle within the sediment. computing trophic indices. However, Marine mud and sand act as a sink for many assumptions are made in many pollutants which then become constructing these indices and multiple bioavailable (it is this bio-availablility feeding behaviour is often not taken rather than absolute concentration into account. A certain degree of which makes the use of the biological trophic information can be gained from components of the ecosystem so meiofauna by using the classification essential in monitoring and effects of nematode feeding types based on studies). With meiofauna the most buccal armature (see Giere, 1993 and sensitive life-stages, the larvae, are references therein). Otherwise, if exposed to this hostile environment, individuals are regarded simply as making meiofauna communities “bits” of information for use in potentially more responsive to statistical analyses (i.e. their pollution events. Most macrofaunal ecosystem function is disregarded), larvae develop in the plankton and are then there is no theoretical difference thus not so exposed. The sedentary between using meiofauna or habit of meiofauna also means that macrofauna sample data, as long as effects are localised. In contrast, samples are taken in the appropriate changes in macrofauna communities way. may be due to “supply-side” events, removed from and unrelated to the Remote sampling for deep water area under consideration. Thus meiofauna is technically difficult and meiofauna show high spatial expensive. High concentrations of resolution. meiofauna inhabit the flocculent layer that is nearly always dispersed by the There are other practical advantages. bow wave of conventional sampling Small size means that smaller gear. No doubt this undesirable effect samples are required with obvious is not confined to meiofauna, but it is implications in the field and for sample assumed that these animals are storage. This can allow a greater disproportionately affected. The degree of replication and statistical inefficiency of macrofauna sampling robustness. Harpacticoids and and sample processing is often nematodes are often relatively overlooked or misunderstood. straightforward to culture and so can

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Meiofauna distribution is dynamic and When is meiofauna sampling most patchy on small (cm) scales. Extra useful? care is needed in designing the Ecosystems are complex and appropriate sampling strategy and obtaining reliable information from setting the correct level of replication. natural populations (by sampling) is Sampling theory still applies but the notoriously difficult. Meiofauna different spatial scales should be taken sampling does not provide a “magic into account. bullet” capable of solving these problems. The weight of current and Identifying meiofauna to species level historic macro-invertebrate research is still the domain of specialists. ensures that the use of meiofauna will However, new identification books and not become prevalent. Nor should this practical manuals are available. The be so. Carefully planned and nematodes can be identified to genus executed surveys using macrofaunal using three Linnean Society Synopses invertebrates are just as valid a way of (Platt & Warwick, 1983, 1988, detecting or tracking environmental Warwick, Platt & Somerfield, 1998) change as similarly diligently and web- and CD-based material from undertaken meiofauna surveys. the Darwin Nematode Project However, there are certain situations (www.pml.ac.uk/nematode/index.html). where high quality samples can be The harpacticoids can be taken to taken in shallow water (by diver or family (and often genus) using another remote coring device) or in intertidal book in the Linnean Society series: habitats, when the use of meiofauna Huys et al (1996). Practical sample should be considered. processing methods may be found in Higgins & Thiel (1988), Giere (1993) One such situation has been alluded and Somerfield and Warwick (1996). to already. This is the case where With these publications, a few frequent monitoring is required to inexpensive items of equipment and provide an early warning system, for some patience it should be possible to example, around a small outfall. In identify the main taxa and produce some such cases the deployment of reference material from a survey in artificial substrata (pan scourers, for any marine biology laboratory. Even if example) may be possible, a taxa cannot be named they can still be technique used by freshwater used for statistical analysis as long as ecologists. This will standardize identification is consistent throughout sampling units, control for differences the study. Nematode and harpacticoid in habitat structure and speed up the experts may then be approached to sampling process. A pilot study will be identify the reference specimens if needed to determine colonization rates necessary. but meiofauna usually colonize new habitats very quickly (Atilla & Fleeger, Further time-saving may be achieved 2000). by subsampling (imperative when meiofauna is abundant). Rapid and Large surveys with many sampling efficient extraction from the sediment sites in areas where meiofauna is can be achieved by flotation methods abundant (e.g. intertidal mudflats) may with centrifugation and sorting (to benefit from targeting the meiobenthos major taxa) is a non-specialist task. in terms of sampling logistics, Some research has shown that transport, storage and the reduced analyses at genus or family level are volume of formalin required. just as sensitive as those conducted at species level. Such taxonomic Finally, in two recent surveys “minimalism” may also be considered undertaken on the south and east as a time-saving device but should be coast of England very few macro- adopted with care. invertebrates were encountered and

PMNHS Newsletter No. 10 April 2002 25 little information was obtained from the sediments. Springer Verlag, Berlin samples. One of these sites, an area 328 pp. of sand swept by strong tidal currents, was teeming with interstitial Higgins, R. P. & Thiel, H. (1988). nematodes and copepods. Other than Introduction to the study of meiofauna those samples quickly investigated out Smithsonian Institution Press, of idle ecological curiosity, all of this Washington DC 488 pp. “information” was carefully washed down the sink. In another study beach Huys, R., Gee, J. M., Moore, C. G. & samples processed for macrofauna Hamond, R. (1996). Marine and produced a few nereid polychaetes. In brackish water harpacticoid copepods. this case, however, contingency was Part 1 Field Studies Council, built into the sampling plan and Shrewsbury Linnean Society meiofaunal animals were extracted so Synopses of the British Fauna (NS) that at least some information could be No. 51 352 pp. gleaned from the survey. In such cases, when it is suspected that Kennedy, A. D. Jacoby, C. A. (1999). macro-invertebrate populations are Biological indicators of marine impoverished, it is always advisable to environmental health: meiofauna - a take meiofauna samples as a potential neglected benthic component? Env. insurance strategy. Mon. Ass. 54: 47 - 68

The few examples here have Platt, H. M. Warwick, R. M. (1983). highlighted the fact that meiofauna can Freeliving marine nematodes. Part I. be of great practical value in British Enoplids Cambridge University environmental monitoring and Press, Cambridge for the Linnean assessment. This has been Society. Linnean Society Synopses of demonstrated many times in academic the British Fauna (NS) No. 28 307 pp. studies. Now that practical methodologies and identification keys Platt, H. M. Warwick, R. M. (1988). are readily available, the meiofauna Freeliving marine nematodes. Part II. should no longer be regarded in the British Chromadorids E. J. Brill/Dr. W. environmental industry as a collection Backhuys for the Linnean Society, of esoteric animals of interest only to London. Linnean Society Synopses of the specialist zoologist. These diverse the British Fauna (NS) No. 38 502 pp. and abundant assemblages can be used with good effect as an important Somerfield, P. J. Warwick, R. M. means of studying and monitoring (1996). Meiofauna in marine pollution environmental change. monitoring programmes. A laboratory manual Ministry of Agriculture, References Fisheries and Food. Directorate of Atilla, N., Fleeger, J. W. (2000). Fisheries Research, Lowestoft 71 pp. Meiofaunal colonization of artificial substrates in an estuarine embayment Warwick, R. M. Platt, H. M. Mar. Ecol. PSZN1 21: 69 - 83 Somerfield, P. (1998). Freeliving marine nematodes. Part III Coull, B. C. Chandler, G. T. (1992). Monhysterids. Field Studies Council, Pollution and meiofauna: Field, Preston Montford. Linnean Society laboratory, and mesocosm studies Synopses of the British Fauna (NS) Oceanogr. Mar. Biol. Ann. Rev. 30:191 No. 53. 296 pp. - 271

Giere, O. (1993) Meiobenthology. The microscopic fauna in aquatic

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The aim of this study is to identify Long-term changes in the characteristic inter-annual time series zooplankton community of the of the North Sea zooplankton North Sea community from CPR data, and then to interpret the shapes of these characteristic time series in terms of climatic signals and particular Richardson, A.J., Lindley, J.A. and taxonomic traits. A technique that has Edwards, M. been applied fruitfully to extracting interpretable patterns from a variety of The Sir Alister Hardy Foundation for data types is the Self-Organising Map Ocean Science (SAHFOS), The (SOM, Kohonen 1997). The SOM is a Laboratory, Citadel Hill, Plymouth, type of unsupervised artificial neural England, PL1 2PB network adept at pattern recognition Email address: [email protected] and classification. There have been over 4300 published papers based on Introduction the SOM, in a wide variety of areas The marine ecosystem of the North including assessing beer quality, Sea suffers considerable human identification of breast cancer, impact from over-fishing, analysing insect courtship songs, eutrophication and possible climate predicting bankruptcies, and in speech change. To assess these human- and fingerprint recognition (see Kaski induced impacts, plankton, which are et al. 1998). This study represents a at the base of the marine food web, preliminary analysis of the suitability of can be used as a biological indicator of the SOM technique to extract the state of the marine ecosystem. interpretable patterns from plankton time series data. The most comprehensive data set on plankton in the North Sea is from the Methods Continuous Plankton Recorder (CPR) Each CPR sample, equivalent to ~10 survey. This survey has provided a nautical miles (18.5 km) of tow and measure of the state of the plankton approximately 3 m3 of water filtered, is community throughout the North analysed under a microscope using a Atlantic on a monthly basis for more standard procedure (Warner and than 50 years. The CPR is an Hays, 1994). The area of interest for automatic plankton sampler towed this study was the central North Sea, behind merchant ships at about 7 m as this was the most comprehensively depth. It filters plankton from the water sampled area during the period 1958- on a constantly moving band of silk. 2000 when the taxonomic resolution of Using this simple but highly cost- the zooplankton counting has effective device, over four million remained unchanged. Only nautical miles have been towed by zooplankton taxa that were present in volunteer ships, collecting nearly more than 50% of the months were 200,000 plankton samples from all included, amounting to 31 taxa. over the North Atlantic. Approximately 400 phyto- and zooplankton taxa are To focus the analysis on long-term counted within each sample. Thus the trends, the SOM analysis was CPR survey is the single largest performed on annual abundances. plankton data set in the world, Annual means were derived by simply providing a unique indicator of averaging the monthly values for each oceanic- and decadal-scale trends in year, as there were no missing months distribution and abundance of plankton in the data set. Prior to the SOM to address global change and analysis, the time series for each ecosystem health issues. taxon was standardised by subtracting its mean and dividing by its standard

PMNHS Newsletter No. 10 April 2002 27 deviation. This allowed the shape of function, with the period to 1981 the time series to be compared, relatively low, and the following years despite the more than three orders of relatively high. The top left series also magnitude difference in the annual has a similar step functions, although abundances between some taxa. there is a massive peak in 1990. The two middle patterns have very low The SOM was performed using the abundance from 1979-1981: most SOM_Pak software Version 3.1 other patterns also reflect this (Kohonen et al., 1995), which is phenomenon, although to a smaller produced by, and freely available from extent. The top right time series has a the Neural Network Research Centre high abundance in the early 1960s, at the Helsinki University of and then flattens out from the late Technology. Input data for the SOM 1960s. The bottom right corner has a consisted of a matrix of 43 (years) very steep negative slope, with an columns by 32 (taxa) rows. A SOM increase in abundance in 1978. with a rectangular topology was used. It was randomly initialised and a two- To relate the characteristic time series step training process was used. The to hydrometeorological events, we first training captures broad-scale have superimposed the timing of the patterns: it had a length of 1000 low salinity (cool temperature) event of iterations, a learning rate of 0.2, and a 1978-1980 and the high salinity event radius of 3. The second training (warm temperature) in 1988-1990 (Fig. reflects finer scale patterns: it had a 2). It appears that the characteristic training length of 10000, a learning abundance series were strongly rate of 0.1, and a radius of 0. affected by these events. It is noteworthy that the low salinity event Results and Discussion led to a decrease in the abundance of A 3x2 SOM of the 31 zooplankton time many taxa, whereas the high salinity series, displaying the characteristic event led to an increase in abundance. time series identified, is shown in Fig. By combining information from Figs 1 1. Taxa that correspond to each and 2, it is possible to identify pattern are also included. Despite the particular taxa that were affected by considerable inter-annual variation in the anomalous hydrometeorological the original time series, the SOM has events. Taxa that had a large decline identified real patterns in the time in abundance during the cold period of series of the various taxa, as the low salinity inflow in 1978-1980 were superimposed raw data series that Centropages typicus, cyphonautes correspond to each pattern confirms. larvae, harpacticoids, hyperiids, Acartia spp., cumaceans, Evadne Patterns are spread in a continuum spp., lamellibranch larvae and mysids, across the two-dimensional space. On and to a lesser extent Calanus the left side of the SOM, patterns are helgolandicus, Candacia armata, generally showing an increase in decapod larvae, echinoderm larvae abundance, on the right side the and post-larvae, fish larvae and patterns are generally decreasing, and Tomopteris. The taxa that had a in the centre the patterns show less of massive increase in abundance a trend. Superimposed on this general corresponding to the warm period of pattern is a considerable amount of high salinity inflow in 1988-1990 were inter-annual variation. A common Calanus helgolandicus, Candacia feature in the patterns in the top row of armata, decapod larvae, echinoderm the SOM and in the bottom right larvae and post-larvae, fish larvae and corner, is a marked peak in Tomopteris; others that were affected abundance around 1989-1990. to a moderate extent were Looking at the SOM in more detail, the Centropages typicus, cyphonautes bottom left series shows a step larvae, harpacticoids and hyperiids;

PMNHS Newsletter No. 10 April 2002 28 those that were affected to a smaller environmental stressors in action? extent were chaetognaths, fish eggs, More research is needed before this gamariids, Limacina spp., Metridia question can be answered. lucens, Calanus finmarchicus, Euphausiids, Oithona spp., This study presents a fruitful and new Para/Pseudocalanus spp. approach of using pattern recognition Pseudocalanus spp. to identify characteristic abundance time series of the zooplankton To identify whether taxa that community, then to link these to correspond to a particular pattern have hydrometeorological events and traits common traits, we have compiled a list of taxa. The current study shows that of some important traits of each taxon many taxa in the North Sea are (Table 1) and superimposed these on affected by hydrometeorologic the SOM output (Fig. 3). These traits conditions, and suggests that taxa with include their permanence in the r-selected traits are doing better than plankton (meroplanktonic or their K-selected counterparts. holoplanktonic), their egg-laying strategy (broadcast or egg-carrying) Acknowledgements and whether they produce diapausal CPR data used in this study are freely eggs. Although there is considerable available to all researchers (see mixing of different traits in the SOM http://www.npm.ac.uk/sahfos/sahfos.ht groups, there are some interesting ml). The authors are grateful to all past general tendencies. For instance, most and present members and supporters taxa on the right side of the SOM are of the CPR survey, especially the holoplanktonic (H), with a greater shipping industry that voluntarily tows proportion of meroplanktonic (M) taxa CPRs on regular routes. The CPR on the left side. In terms of spawning survey has been recently funded by a behaviour, there are more broadcast consortium comprising IOC, and spawners (B) on the left side than are agencies from Canada, The Faroes, on the right side. And in taxa that have France, Iceland, Ireland, Netherlands, diapausal eggs (T), there are none (F) Portugal, UK (DEFRA and NERC) and on the right and many on the left side. USA.

It is noteworthy that taxa that have References some or all of the traits of being Kaski, S., Kangas, J. and Kohonen, T. meroplanktonic, a broadcast spawner, (1998) Bibliography of Self-Organizing and having diapausal eggs are Map (SOM) Papers: 1981-1997, generally increasing in abundance Neural Computing Surveys, 1: 102- (more common on the left side of the 350. Available in electronic form at SOM) (also see Lindley and Batten http://www.icsi.berkeley.edu/~jagota/N 2002). These traits correspond to an r- CS/. selected strategy, well adapted to a Kohonen,T., Hynninen,J., Kangas,J. variable or stressed environment. By and Laaksonen,L. (1995) SOM Pak contrast, taxa with traits of being Version 3.1. holoplanktonic, egg sac spawners and Kohonen, T. (1997) Self-organizing not having diapausal eggs have maps. Springer, Berlin. decreased in abundance (right side of Lindley, J.A. and S.D. Batten (2002) SOM). These traits correspond to a Long-term variability in the diversity of more K-selected strategy North Sea zooplankton. J. Mar. Biol. (holoplanktonic, egg sac spawners Ass. U.K. 82: 31-40. and no diapausal eggs), more suitable Warner, A.J. and G.C. Hays (1994) for a less variable or more less- Sampling by the Continuous Plankton stressed environment. Has there been Recorder survey. Prog. Oceanogr. 34: an increase in the variability of the 237-256. environment in the North Sea, or are

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Table 1. Traits and preferences for each taxon used in the analysis. These include their permanence in the plankton (holoplanktonic (H) or meroplanktonic (M)), their egg-laying strategy (broadcast (B) spawner or egg-sac spawners (E)), and whether they produces resting eggs (true (T) or false (F)).

Taxa Permanence Egg Laying Resting in Plankton Eggs Acartia spp. H B T Calanus finmarcicus H B F Calanus helgolandicus H B F Candacia armata H B T Centropages hamatus H B T Centropages typicus H B T Chaetognatha H B F Cirripede larvae M E T Cumacea M E F Cyphonautes larvae M E F Decapoda larvae M E F/T Echinoderm larvae M B F Echinoderm post-larvae M B F Euphausiacea H B F Evadne spp. H E T Fish larvae M B F Gammaridea M E F Harpacticoida H E F Hyperiidea H E F Lamellibranch larvae M E F Larvacea H B F Limacina retroversa H E F Metridia lucens H B F Mysidacea H E F Oithona spp. H E F Para-pseudocalanus spp. H E F Podon spp. H E T Polychaeta larvae M B F Pseudocalanus elongatus H E F Temora longicornis H B T Tomopteris spp. H B F

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Fig. 1. A 3x2 SOM of the 31 zooplankton taxa in the North Sea, showing the characteristic time series identified (dark line) and the raw data series for each taxon superimposed (dotted line). Taxa corresponding to each raw data series is also shown.

Fig. 2. The low salinity (1978-1980) and high salinity (1988-1990) events superimposed on the characteristic time series identified by the SOM.

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Fig. 3. Traits of taxa that mapped to each SOM pattern. Codes for variables are: These include their permanence in the plankton (holoplanktonic (H) or meroplanktonic (M)), their egg-laying strategy (broadcast (B) spawner or egg-sac spawners (E)), and whether they produces resting eggs (true (T) or false (F)). The order of the taxonomic traits for each SOM pattern is the same as that of the taxonomic names in Fig. 1.

Broad scale mapping in the Forth:The Firth of Forth Spatial Project

Dr Richard Park

Senior Marine Biologist, SEPA, Clearwater Scotland, Heriot-Watt University, House, Heriot Watt Research Park, Scottish Natural Heritage and Scottish Avenue North, Riccarton, Edinburgh EH14 Environment Protection Agency in the 4AP [email protected] FFSS and briefly discuss the relevance of this review to the Urban

WWTD and Water Framework Abstract only Directive. The presentation will include The Firth of Forth Spatial Study examples of broad scale mapping of (FFSS) aims to review the current sediment type and benthic habitats status of soft sediment benthos in the within the inner firth. Forth. This short presentation will describe the collaborative interests and roles of National Museums of

PMNHS Newsletter No. 10 April 2002 32

parallel to the Arms Race, came the Challenges in Marine Space Race. Kennedy’s commitment Biodiversity: The Global View to put a man on the moon within 10 years assured NASA of limitless funds for space exploration. But it was the Voyager missions, the exploration of the outer reaches of the solar system by small smart probes that really A. D. McIntyre caught my imagination.

63 Hamilton Place, Aberdeen AB15 5BW In a way, though, the explorers of [email protected] space had it easy. The missile programme had solved the launcher Abstract only problems, so escaping the Earth’s This talk considers how the study of gravity was eminently feasible, if marine biodiversity has developed expensive. Once in space the rest was over the years, looking first at the early straightforward – limitless supplies of days when the work was not graced energy from solar panels, or your with a formal name. The study went personal chunk of plutonium as in the through a difficult period when Voyager craft; no motion resistance, identifying organisms was not seen as no corrosion, and so on. But the a relevant science, but the importance greatest boon of space was its of biodiversity is now globally transparency to the electromagnetic recognised, and its study is spectrum, allowing unfettered optical increasingly supported. Its impact on and radar imaging of planetary the global scene is noted, and a surfaces and, more significantly, ease relevant emerging research of radio-communication with the programme is briefly discussed. spacecraft.

However, my university education in physics, coupled with several summers exploring in Greenland, led Marine technology: challenges of the last frontier not to space science but to the Scott Polar Research Institute. There I joined a team using airborne radars to map the Antarctic continent, hidden under several kilometres of ice. D. Meldrum Unfortunately ice is not as transparent as space: powers that could bounce Leader, Technology Development signals off the moon are needed to Scottish Association for Marine Science, generate detectable echoes through 5 Dunstaffnage Marine Laboratory, Dunbeg, km of polar ice. But at least radio Oban PA37 1QA waves do penetrate ice. And we did Scotland [email protected] find the unexpected – Lake Vostok, deep in the heart of the continent, First published as THES 'Cutting Edge' under more than 4 km of ice. feature, 7 Sept 2001, reproduced by permission of the Times Higher When this programme wound down I Education Supplement, moved back to Dunstaffnage in www.thesis.co.uk Scotland to study a natural medium that is, for all practical purposes, totally I grew up in cold war Scotland, where opaque to radio waves – the sea. regular reports of missile and bomb Because of this simple fact the bottom tests imbued our daily lives with a of the oceans is much less well distinct sense of unease. Then, in mapped than, say, the far side of the

PMNHS Newsletter No. 10 April 2002 33 moon, or the surface of Mars. Satellite- universities. Instrument pods, borne instruments, which have done connected to the network via sea-floor so much to advance our knowledge of junction boxes, will both deliver data the Earth’s surface and atmosphere, and receive commands and energy. are powerless to see beneath the AUVs will now be able to dock surface of the ocean. Seabed mappers routinely to report their findings and have to rely on acoustic instruments to recharge their batteries. The spirit of build up images of the ocean floor, and exploration which has led to such these instruments have to be within amazing discoveries as the volcanic the water. Unlike space, water resists activity on Jupiter’s moon Io, and Lake the motion of objects through it: Vostok in the Antarctic, is about to be building the oceanographic equivalent fostered once more in the oceans. of a satellite – an autonomous And, connected to the Internet, what underwater vehicle (AUV) – faces the new wonders will our last frontier immediate problem of supplying the reveal – and to whom? vehicle with enough energy to navigate. And, unlike the days of Voyager, small nuclear reactors are no longer popular. Practical developments in numerical analysis techniques Water friction can, however, be turned used for marine biological to advantage: the oceans move, and it surveys is possible to hitch a free, if largely unpredictable, ride on the back of ocean currents. At Dunstaffnage I have become closely involved in Annette Woodham exploiting this free ride, and in recycling new technologies such as ERT (Scotland) Ltd, Research Avenue 1, GPS for the detailed study of ocean Heriot-Watt University, Edinburgh EH14 currents. We have deployed drifting 4AP instruments in many areas, including Abstract the polar ice packs, where they have Marine biological surveys usually yielded new data for the climate generate complex datasets, change debate. Elsewhere, the representing the presence and profiling floats of the international Argo abundance of numerous species at programme drift for days on deep different sampling locations and currents, surfacing periodically to sometimes also at different times. download thermal data that will help us Various numerical methods have been understand the interplay between the developed to help summarise the data oceanic heat reservoir and climate and to identify trends in spatial change. distribution or changes over time. The

last 25 years have seen enormous This still leaves the ocean floor largely developments in the availability of unmapped: speculative computers and vast increases in reconnaissance – exploration in the computational power. The talk true sense – is largely ruled out. examines how the numerical methods However, change is afoot, spurred by available for analysing survey data events such as the chance discovery have changed over the this period, of the biological communities thriving and how this may have influenced our around hydrothermal vents. Plans are working practices and our well advanced in many countries for interpretation of survey data. Practical the establishment of ocean examples are drawn from experiences observatories linked by sea-floor in our own laboratory, which was cables - an Internet of the deep ocean, founded in 1976. In more general open to primary schools as well as

PMNHS Newsletter No. 10 April 2002 34 terms, modern program packages Objectives of data analysis have made complex multivariate A variety of numerical methods have techniques widely available and easy been developed to help summarise to use, but is there a downside to this? large datasets and make them easier And what aspects of data analysis to interpret. These techniques help to remain fundamentally the same identify trends in spatial distribution or despite the developments in computer changes in community composition technology? over time. We have used them in particular to detect impacts from Introduction possible sources of pollution, as part The aim of the talk is to look at how of monitoring programmes. These the considerable developments in programmes usually involve computer technology over the last 25 conducting baseline and post- years have affected the way we operational surveys, with samples conduct numerical analysis of marine taken from a number of stations that biological survey data. I thought it may be affected and from reference would be interesting to look at how stations. Numerical analysis these changes may have affected the techniques are also used in pure way we work, for example in terms of research aimed at describing how we collect data and how we communities and their distributions. interpret the results of surveys. Emphasis is on the practical aspects Types of numerical analysis of conducting the analyses, rather than The numerical techniques used fall on theoretical developments. I am into three general types: univariate drawing on the experiences in our own methods, graphical methods and laboratory at ERT, founded in the mid- multivariate methods. Univariate 1970s at the Institute of Offshore methods involve looking at simple Engineering (IOE) at Heriot-Watt community statistics such as the University. The talk was put together number of species and the number of by speaking to people who analysed individuals, and a variety of diversity data there at different times in the last and evenness indices which are 25 years, and is illustrated with calculated from the numbers of examples of how we actually worked species and individuals and the and the problems tackled. I was distribution of individuals among the analysing survey data myself mostly species. Univariate statistics are between 1990 and 1996 and saw calculated for each sample or station considerable changes even over that separately, and can then be used to relatively short period of time. Of compare stations with each other. The course, other laboratories will have various graphical methods, such as done things differently, but I think the rarefaction curves, log normal general evolution of trends is quite distributions and ABC curves, are typical. different ways of plotting basic univariate statistics onto graphs so What kind of data? that their shapes can be compared for The kind of data I am referring to are different stations. those generated from the processing of grab or core samples taken Multivariate methods use the whole subtidally or intertidally. The fauna in dataset from a survey or series of each sample is extracted, identified surveys, and compare each sample and counted, and sometimes weighed. with all the others. They take into This results in a large number of account the identities of the species at records of abundances of different each station, as well as their species at different places, and abundances. Such methods include: sometimes also at different times. cluster analyses, which group stations according to their faunal similarity,

PMNHS Newsletter No. 10 April 2002 35 represented by a dendrogram (fork curves. Later on, Simpson’s (D) and diagram); gradient analyses, which Brillouin’s (Hb) diversity indices and detect linear gradients in the faunal Heip’s evenness measure (Eh) were data; and multidimensional scaling, added, and Hurlbert’s rarefaction which maps stations in a way that curves replaced Sander’s. reflects their faunal similarity. Multivariate statistical techniques were Timeline not used routinely for marine It emerged that the changes in monitoring, and the reports from that practices could best be presented by time instead provide relatively detailed dividing the period since 1976 roughly discussions on the distributions, into four time periods that reflect the abundances and biology of individual most significant developments. The species in the community. Univariate practicalities of data analysis during data were calculated and compared the different periods are outlined for each phylum in addition to the below. The divisions are not distinct whole dataset. but convenient. The changes occurred to various aspects of the The reports were typed on typewriters process: data entry and management; of course, including the species lists data analysis; and output and which caused considerable problems presentation of results. These aspects with spelling. Mistakes had to be are considered for each time period. corrected by retyping the whole page, but each time this was done other Approximately 1976 to 1982 mistakes occurred, so pages were At this time there were no PCs, but corrected until the number of mistakes access could be gained to mainframe was constant. computers at the universities. Early computers became available in The survey data were collated by the late 1970s, and IOE used BBCs transcribing all of the species- and a ‘Pet’. These had 16 or 32 abundance records from the individual Kbytes RAM and a clock speed of 4 site sheets onto large hand-written Khz, and there was no hard disk. This spreadsheets. Firstly, the names of all leads into the next period. the species recorded were listed in the first column, using the contents list in A BBC computer Hartmann-Schröder (1971) as a guide to taxonomic order for the many polychaetes. A system of squares was used to record the abundances of each species in each sample (replicate) at each station. The numbers were checked by manually adding up columns in both directions. Sue Hamilton worked for IOE at the time and used to do this on her kitchen table.

Univariate statistics were calculated by hand. Examples used in early surveys Photograph: Philippa Kingston included Margalef’s diversity index (d), the Shannon-Wiener information Approximately 1983 to 1987 IBM PCs came in and started to be function (Hs) and Pielou’s evenness measure (J). Graphical techniques used for some univariate analyses, but used included Preston’s log normal multivariate analysis was carried out distributions and Sander’s rarefaction on a mainframe computer.

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The data were typed directly into the PCs as data files in specified formats. We used the program ‘Stirling 3’ and later versions (Moore, 1983), which provided a list of the ten most abundant species at each station, together with a range of diversity indices (Hs, D and Hb) and evenness measures (Eh and J), and the co- ordinates for calculating Hurlbert’s Photograph: Paul Kingston rarefaction curves. The results were presented only as a hard copy via the The results from the analyses had to printer onto computer paper in a be plotted by hand, and figures for continuous roll. The process was very reports were drawn on tracing paper slow - we would wait with anticipation using Rotring pens and Letrasett. for the next co-ordinate for a Hurlbert’s curve to be generated. Consequently the program was often left to print Approximately 1988 to 1993 overnight, and then by the morning the Faster PCs became available over this paper would have jammed. period, so that we could now carry out all analyses in-house. All of our We started to use multivariate software was DOS-based, and there analyses routinely during this period, was much less standardisation of and these were carried out using a general software, such as mainframe computer, initially at spreadsheets and word processors, Aberdeen University and later at than there is today. Heriot-Watt. The first program used was ‘clustan l. c.’ (Wishart, 1978), for A key development in the laboratory cluster analysis, and was frequently was the creation of a system for carried out on binary data (species entering data easily and accurately. presence and absence). Later, the IOE commissioned Dr Colin Moore of FORTRAN IV versions of the Cornell Heriot-Watt University to write a Ecology Program series were used, program for data entry. This resulted particularly a type of gradient analysis in WORMS, a data entry and called detrended correspondence management program (Moore, 1989, analysis or Decorana (Hill, 1973). The unpublished). Based on the database data had to be input in FORTRAN DB3+, WORMS sorts entries into format, which was achieved by taxonomic order and provides output laboriously writing out coding cards. files for numerical analysis. Use of Accurate placement of all of the WORMS involved developing a species abundance data (and zeros) in species coding system as part of our the grid boxes was essential to the in-house species directory. Each successful running of the programs. species was allocated an 8-character The coding cards were in turn used to alphanumeric code and a brief code make up punch cards; one punch card consisting of a 3X3 couplet. The for each species, ie each column of nomenclature and taxonomic order data. Initially we supplied data on were based on the Directory of British coding cards, but later had access to a Marine Fauna and Flora (Howson, punch card machine ourselves. This 1987). A rather dog-eared photocopy was the size of a desk and resulted in of this had all of our species code a box of punch cards maybe 60-90 cm numbers written in it, so that we would long for a single survey. know how to code any additional species we found. A punch card used for data input to mainframe computer

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Later versions of STIRLING (why did well, but Excel was easy to use with we start using upper case for the new software packages. However, programs?) were used for univariate we still use our species directory analyses, while the program package database, with its 8-character coding PATN (Belbin, 1988) was used for system and ‘brief’ codes, to sort multivariate analyses. The latter were species into taxonomic order, provide dominated by cluster analysis and correct spelling and authorities, and DECORANA. Multidimensional help us manage our species reference scaling became feasible as PCs collection. became more powerful, and later on in this period we started using the For univariate analyses, a program program MDS, amongst others, in an DIVCALC was written for us by Rob early version of the PRIMER package Rolph, to provide output exactly as we developed at the Plymouth Marine wanted it. Some projects, such as Laboratory. surveys around fish farm cages and sewage outfalls, required use of the The main impression I have from this Infaunal Trophic Index, which is time is that no software was calculated for each station based on compatible with anything else. Each the feeding types of the animals there, program required data to be input in its so Rob also wrote ITICALC. own particular format. We had programs specially written to convert More powerful computers mean that data into different forms for different multivariate analyses can now be packages. The conversion of the data carried out much faster than before. for all the different programs, and for This is especially noticeable with MDS, the tables we needed for our reports, in which the whole computation is was quite complicated, and one of my repeated several times in order to jobs was to write the laboratory achieve the plot that best represents computer manual to remind us all what the data. When I first started using to do. There were also limitations on PRIMER I chose 5-10 repeats the amount of data that could be because this could be completed analysed at once; one ten-year within an hour or less. Now we dataset had to be split in half for some routinely do 40 and it takes seconds. of the techniques we wanted to use. The packages we use most commonly now are PRIMER for Windows; MVSP The quality of the output from PATN (Multivariate Statistical Package) as a was not suitable for reports, and the more modern package for co-ordinates from DECORANA and DECORANA and other programs; and cluster analysis were plotted PC-Ord. afterwards from the results files generated. By this time we had Software generally has become more invested in Apple Macs for our desk- compatible in recent years. PRIMER top publishing, which only added to output was always neat but we used to our incompatibility problems. redraw figures on the Macs. Now, graphics output is easily incorporated Approximately 1994 to present into reports and we have much more PCs have become increasingly flexibility in presentation. Also we now powerful, with 386s, 486s and then use PCs for report production. Pentium processors. We eventually started using Microsoft Windows in the Discussion laboratory, with some resistance. The developments in computer technology over the last 25 years that Around 1996, we started using have affected the way we analyse and Microsoft Excel spreadsheets to enter present marine biological survey data data. WORMS had served us very include both hardware developments

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(the availability of PCs and then vast What hasn’t changed in the last 25 increases in computer power) and years? Despite all the benefits of software developments (both general technological advances mentioned and specialist software). above, many aspects of data analysis remain fundamentally the same. The These have made numerical analyses same univariate statistics tend to be easier to carry out and more widely used, for example, and cluster used. Complex multivariate analysis and DECORANA have stood techniques have become much more the test of time. user-friendly and a wide range of techniques is now readily accessible. References Analysis is fast, allowing us to carry Belbin, L. (1988) PATN: Pattern out a number of different tests to analysis package. CSIRO Division of ‘explore’ a dataset. We can handle Wildlife and Rangeland Research. bigger data sets, combining data collected over long-term studies and Hartmann-Schröder, G. (1971) analysing them together. We also Annelida, Borstenwürmer, Polychaeta. have tremendous flexibility in the way Die Tierwelt Deutschlands, 58: 1-594. we present survey findings. These are all positive aspects, but are there any Hill, M.O. (1973) Reciprocal averaging: downsides to all this progress? an eigenvector method of ordination. Journal of Ecology, 61: 237-251. Can it be too easy? I have heard examples of students using Howson, C.M. (1987) Directory of the multivariate software packages without British Marine Fauna and Flora. A really understanding what they are coded checklist of the marine fauna doing with their data, and and flora of the British Isles and its misinterpreting the output. This may surrounding seas. Marine always have been the case, but the Conservation Society, 471 pp. earlier packages demanded a lot more study before you could operate them, Moore, C.G. (1983) Stirling 4. A Basic and you were more in touch with what program for the investigation of was happening to the data. There species diversity. Wat. Pollut. Control, may also be an impression now that 82: 102-106. the techniques can do more than they can, ie that they can provide all the Wishart, D. (1978) Clustan User answers, rather than just being tools. Manual (3rd edition). Inter- University/Research Councils Series, Also, I wonder if there is a tendency to Report No 47, 175pp. adopt methods that are readily available, at the expense of other Acknowledgements methods. A certain amount of I am very grateful to the following for conformity is necessary in terms of useful discussions and for providing environmental regulation, but we must anecdotes, records and images from avoid over-standardization and think the different times: Sue Hamilton, Paul before we press the default button. Kingston and Colin Moore from Heriot- Another consideration is whether we Watt University; Peter Campbell, are putting too much emphasis on Claire Dalgleish, Iain Dixon and multivariate routines generally, at the Jonathon Hunt from ERT (Scotland) expense of looking at the species list. Ltd. Thank you also to David Philip for Early survey reports focused more on helping to prepare the presentation. individual species and their distributions, and a wider range of univariate statistics were used.

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